Full text of "Reports and papers on botany"

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OF CALIFORNIA

DAVIS

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INSTITUTED MDCCCXLIV.

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LONDON.

MDCCCXLIX.

BflHY

REPORTS AND PAPERS

BOTANY.

CONSISTING OF

I, MOHL ON THE STRUCTURE OF THE PALM-STEM.
II. NAGELI ON VEGETABLE CELLS.

III. NAGELI ON THE UTRICULAR STRUCTURES IN THE
CONTENTS OF CELLS.

IV. LINK'S REPORT ON PHYSIOLOGICAL BOTANY FOR 1844-45.
V.GRISE BACH'S REPORT ON GEOGRAPHICAL BOTANY FOR 1844.

VI. GRISEBACH'S REPORT ON GEOGRAPHICAL AND
SYSTEMATIC BOTANY FOR 1845.

EDITED BY

ARTHUR HENFREY, F.L.S.

&c. &c.

LONDON:
PRINTED FOR THE RAY SOCIETY.

MDCCCXLIX.

LIBRARY

UNIVERSITY OF CALIFORNIA
DAVIS

PRINTED BY C. AND J. ADLARD,

BARTHOLOMEW CLOSE.

CONTENTS.

ON THE STRUCTURE OF THE PALM-STEM.
BY PKOF. H. VON MOHL.

PAGE
Introduction . . ... . . .3

On Palm-Stems . . . .5

Form of the Palm-Stem . . . . . ib.

Course of the Vascular Bundles in the Stem . , 7

Cellular Tissue of the Palm-Stem . . .10

Structure of the Vascular Bundle . . .15

Modifications of the Structure of the Vascular Bundle in

various Palm-Stems . . . .19

Of the Structure of the particular Anatomical Systems of

the Vascular Bundle . . . .20

Comparison of the Palm-stem with the stems of other

Monocotyledons . . . . .30

Comparison of the Vascular Bundle of the Palms with that

of the Dicotyledons . . . . .36

Comparison of the Palm-stem with the Stem of Dicotyledons 39

On the Root of the Palms . . . . .45

Form of the Root . . . . . ib.

Anatomical Examination of the Root . . .46

Appendix ... .51

VI CONTENTS.

ON THE NUCLEI, FORMATION, AND GROWTH OE
VEGETABLE CELLS. BY PROF. C. NAGELI.

PART II.

PAGE

IV. Free Cell-formation .... .95

a. Without Visible Nucleus . . . . ib.

b. With a parietal Nucleus .... 103

c. Of free Cell-formation as a general Law . .111
V. On Cell-formation in general . . . .123

VI. On the Growth of Cells . . .143

ON THE UTRICULAR STRUCTURES IN THE CONTENTS
OF CELLS. BY PROP. C. NAGELI.

Introductory . . . . . . .161

1. Nuclear Utricles, Nuclei . . . . .165

2. Spermatic Utricles . . . . .171

3. Nucleoli . . . . . . . ]72

4. Mucilage- (Protoplasm-) utricles . . . .173

5. Proliferous Utricles . . . .175

6. Colour-utricles . . . . . .176

7. Starch-utricles, Starch-granules . . . .183

8. General Retrospect . . . . .188

REPORT ON PHYSIOLOGICAL BOTANY FOR 1844 AND 1845.
BY PROP. LINK.

General Observations ..... 193

I. Internal Structure of Plants .... 221

II. Stem and Root . . . . . .249

III. Root, Tubers, Prickles, Tendrils, Glands, Stomata . . 264

IV. Leaves . . . . . . . 270

V. Flowers, Fructification . . . . .282

VI. Fruit, Seeds, Germination ..... 29O
VII. Individual Orders and Genera of the Phanerogamia in Refer-
ence to Physiology . . . . .296
VIII. Ferns, Mosses, Lichens, Algae, Fungi . . . 304
IX. Monstrosities ... 312

CONTENTS. Vll

REPORT ON GEOGRAPHICAL BOTANY FOR 1844.
BY PROF. GRISEBACH.

PAGE
Introductory . . . . . . .317

I. Europe . . . 323

II. Asia. , . - 349

III. Africa .... .377

IV. Islands of the Atlantic Ocean . . . .388
V. America 390

VI. Australia and South Sea Islands . . .407

REPORT ON GEOGRAPHICAL AND SYSTEMATIC BOTANY
FOR 1845. BY PROF. GRISEBACH.

A. Botanical Geography . . . . . .418

I. Europe . . . . . .423

II. Asia. . . . 455

III. Africa ... . 457

IV. America . . . . . .458

V. Australia and South Sea Islands .... 469

B, Systematic Botany ...... 474

EXPLANATION OF THE PLATES.

Mohl on the Palm-Stem . . . . .495

Nageli on Vegetable CeUs . . . . . ib.

INDEX to the Physiological Report . 503

Geographical Botany . . . . 507

Systematic Botany . . . . .512

ON THE

STRUCTURE OF THE PALM-STEM.

BY HUGO VON MOHL.

(The Treatise ' de Structura Palmarum' contained in Martius's ' Genera et Species Palmarum.')

TRANSLATKD, WITH THE APPENDIX,

(From the German version, published by the Author, in his ' Vermischte Schriften botanischen
Inhalts' Tubingen, 1845,)

BY AKTHUR HENFREY, F.L.S. ETC.

.C.D. LIBRARY

STRUCTURE OF THE PALM-STEM.

INTRODUCTION.

A minute anatomical examination of the Palms is of
especial importance in regard to the anatomy and phy-
siology of plants, because the characters of the Mono-
cotyledons are most clearly exhibited in them, and they
therefore afford the most favorable means of acquiring
satisfactory ideas of the structure and growth of this great
class of plants. As the earlier phytotomists had devoted
but little care to the examination of Palm-sterns,* these
suddenly acquired very great importance when Daubenton,
in the examination of the Date-palm, believed that he
found the vascular bundles proceeding to the young leaves,
becoming developed in the interior of the stem, surrounded
by the vascular bundles running to the older leaves. This
proposition, forming an epoch in the history of Phytotomy,
first appeared in its full importance when Desfontainesf
showed that, not only in the Date-palm, but in Mono-
cotyledons generally, the wood has the form of scattered
vascular bundles, and that the vascular bundles which run
to the leaves come from the centre of the stem. This

' Vide Grew's Examination of Calamus, Anat. of Plants, p. 104.

f Mem. sur 1'Organisat. d. Monoc. (Mem. de PInstitut National, t. i, 478).
According to a brief notice published by Mirbel (Comptes Rendus, 12 Juin,
L843), the first discovery belongs to Desfontaines, he having already, in his
c Travels in Algiers/ expressed in a few words the idea which lies at the
foundation of his system.

4 THE PALM-STEM.

discovery necessarily excited the greatest attention ; the
doctrine, that the vascular bundles of the Monocotyledons
originate in the centre of the stem, and press the older
bundles outward ; that this process continues until the
older and solidified vascular bundles form a layer at the
circumference of the stem sufficiently firm and hard to
resist the pressure of the younger; that thereupon all
further increase of thickness of the stem must cease, and
that from this results the columnar form of the stem this
doctrine appeared to explain the peculiarities of the growth
of Monocotyledons in a manner so simple and satisfactory,
that it not only passed into all text-books, but was even
used by De Candolle for the systematic division of vascular
plants into Endogens and Exogens. The observations of
Dupetit-Thouars * showed, indeed, that the stem may
grow to an unlimited thickness in many Monocotyledons ;
they were not, however, any more than the later obser-
vations of Mirbel, sufficient to shake the belief in the
correctness of Desfontaines' doctrine, and only gave oc-
casion for the assumption that in some Monocotyledons a
second peripherical growth occurs, in addition to the
central vegetation. One voice, alone, but therefore the
more important, declared against the theory of the central
growth of Monocotyledon s.f Moldenhawer stated that in
the stem of Phcenix dactylifera a line of division occurs,
on both the outside and inside of which liber-bundles
are equally developed; that around those in the latter
situation, spiral vessels are produced subsequently to their
first formation, and they thus become ligneous bundles ;
and that of these ligneous bundles, the inner run to the
older leaves, and the outer to the younger; that con-
sequently, in other words, the date-palm has a periphe-
rical growth. Like so many other admirable remarks of
this exact observer, this proposition was so completely
neglected by other vegetable anatomists, that not one
thought it even worth the trouble of mentioning.

* Premier Essai sur la Vegetation.

f Beitrage zur Anatom. d. Pflanzen, 53.

THE PALM-STEM. 5

ON PALM-STEMS.
Form of the Palm-stem.

The organization of the Stem of Palms undoubtedly
exhibits common characters throughout all species : it is
simple (except in Hyphcene), almost cylindrical, generally
upright, encircled by the scars of the leaves which sur-
round the stem, but without true nodes; fixed to the
earth by slender fibrous roots ; having at the apex a tuft
of leaves, in the axils of which stand the spadices. The
internal structure is, generally speaking, as follows : the
groundwork of the whole stem is an uniform, lax paren-
chyma, in which lie, scattered irregularly, slender vascular
bundles, running apparently parallel with the axis of the
stem; those lying nearest to the circumference of the
stem are mostly thicker, of a more woody consistence,
and placed closer together than those situated in the
interior ; from this circumstance, the stem often possesses
very considerable solidity toward the periphery.

For the convenience of anatomical description, I divide Palm-stems into
some subdivisions, which, however, do not correspond to the systematic
subdivisions based on the modifications of the structure of the flowers and
fruit.

1. The Cane-Vko, (Geonoma-like) palm-stem, Caudex arundinaceus, is thin,
slender, upright, with the nodes tolerably near together, the internodes
obconical, the epidermis smooth, shining, and not decaying from the action
of the atmosphere. These stems are moderately strong, the parenchyma is
simple and close, the fibres lying in the middle of the stem softer, those at
the circumference often pretty hard, the liber-like layer weak. At first sight,
these stems bear much resemblance to the stems of Grasses, especially of
bamboo, to which the yellow colour acquired on drying, and the obconical
form of the internodes, which give the stem an articulated aspect, much con-
tribute ; they are, however, easily distinguished from, the culms and subter-
raneous stems of Grasses, by the absence of a central cavity, and by the
circumstance that the vascular bundles do not form any reticuiarly branched
interlacement at the nodes. This stem occurs in most species of Geonoma,
many species of Bactris, in Hyospathe, Chamtedorhea ; similar forms, but

6 THE PALM-STEM.

constituting the transitions to the other forms of stems, are found in
Desmoneus, Rhapis flabelliformis, and Corypha frigida.

2. The Calamoid stem (Caudex calamosus) resembles the cane-like, but is
distinguished by the extraordinary length. The internodes are from two to
six feet long, thin, apparently cylindrical, but equally obconical ; the surface
smooth, shining as if varnished ; of a stony hardness. The substance is not
firmer at the periphery than in the middle ; the vascular bundles are distri-
buted almost uniformly throughout the whole stem; the woody mass is
moderately hard, exceedingly elastic, and tough ; the external fibrous layer
is very thin ; the stony, hard epidermis splits off in scale-like pieces when the
stem is bent. This form occurs only in Calamus ; the transition to the cane-
like stem is through Desmoneus.

3. The Cylindrical (Mauritia-likz) stem is remarkable for its beautifully
regular uniformity and smooth, round form. The internodes are pretty long
and cylindrical ; the leaf-scars are narrow and do not form knot-like pro-
jections ; the rind thin, not much affected by the action of the atmosphere,
often clothed with spines. The internal structure is very strongly marked ;
almost the whole mass consists of a weak, lax, pith-like parenchyma, in which
lie vascular bundles of herbaceous softness. Firm woody vascular bundles
are only found at the periphery in a narrow circle ; but on account of their
frequently considerable thickness and hardness, they form an almost impene-
trable layer. The external fibrous layer is mostly very thin. This form
occurs in Mauritia (armata, viniferd), (Enocarpus (minor, &c.), Kunthia
(montana), Astrocaryum (vulgare, &c.)

4. Cocos-like stem (Caudex cocoides). This is thick, and somewhat irre-
gularly knotted from the closely approximated, broad leaf-scars, and fre-
quently shaggy with the vascular bundles of the fallen leaves and withered
leaf-sheaths ; often very tall. The vascular bundles are distributed almost
uniformly throughout the whole mass ; those situated near the periphery are
merely a little more closely approximated than the internal, and are rather
thinner than thicker than the latter. The liber-like fibrous layer is very
thick ; the rind thick, irregularly torn, and weathered. The ligneous bundles
are sometimes rather soft, as in Corypha cerifera, but also sometimes very
hard, as in Cocos coronata. On account of the uniform distribution of the
vascular bundles, the stem is nearly as hard in the middle as in the outer
part, and on account of the great number of bundles, it presents considerable
solidity. This form of stem occurs in Cocos, Leopoldinia, Syagrus, Mais,
Corypha ; Ehapis Jlabettiformis and Lepidocaryum gracile form the transition
to the cane-like stems.

5. Stemless Palms. In some the length is so small that the plants seem
to be stemless. Two varieties occur. In the first the stem is abbreviated,
like a bulb. This is not peculiar to any special genus, but occurs in indivi-
dual species of the most varied genera, e. g. in Geonoma acaulis, macrostachys ;
Astrocaryum acaule, campestre ; Diplothemium maritimum, campestre, littorale.

THE PALM-STEM. 7

Isolated species occur with stems sometimes very much abbreviated, and at
other times of tolerable length, e. g. Attalea compta. The second variety
occurs in Sabal here the stem forms a short, creeping rhizome, of a most
remarkable form, its leafy apex lying on the ground, while the hinder ex-
tremity is lifted up by the roots, and projects above the ground.

Note. I had no specimens of this fifth form to examine ; the following
remarks relate, therefore, only to the first four forms of stems mentioned.

Course of the Vascular Bundles in the Stem.

Before I proceed to the microscopic anatomical de-
scription of the stem, it will be necessary to describe the
course of the vascular bundles. It is known that these do
not lie in concentric circles, but are scattered without
definite arrangement throughout the stem. This differ-
ence of the Palms from Dicotyledonous trees is so striking,
that even in ancient times it was regarded as a charac-
teristic peculiarity of Palms.* The course of the vascular
bundle is best traced in stems where the parenchyma has
lost its firmness by decomposition ; in these the individual
bundles may with very little trouble be extricated from a
stem split longitudinally. Stems with a white pith-like
centre are also very well adapted to this investigation.
When, in such a stem, e. g. Kunthia montana, a vascular
bundle is traced from the point of insertion of the leaf
backward, it is found that it runs in a curve (the con-
vexity upward,) to the centre of the stem, then in the
neighbourhood of the centre runs down a certain extent
deep in the stem, but soon again loses the direction
parallel to the axis of the stem, gradually (since at the
same time it is always running down the stem) again
approaching the surface, till it lies beneath the rind, and
there passes down the stem beneath this.

Otis. I have here described the course of the* vascular bundle in the
direction from above downward, because I usually traced them in this

* Theophrast., Hist. Plant., lib. i, cap. ix.

O THE PALM -STEM.

direction in the stem ; but it must not be hence inferred that the vascular
bundle of Palms is perfected in this direction in its formation ; and in the
following pages I shall, according as it may be more conveniently stated,
follow the vascular bundle, in the anatomical description, sometimes from
above downward, and sometimes in the opposite direction.

The course of the vascular bundle is the same in all
Palms, and the only distinctions which present themselves
are a difference of aspect at different points of the course
of the vascular bundle in different species.

In those species, for instance, which, like Kunthia
montana and Manritia aculeata, possess firm woody vas-
cular bundles only at the periphery of the stem, with a
centre composed of soft, herbaceous substance, we find
that all the bundles are thin, soft, and herbaceous from
the point where they enter the leaf, downward to the
centre of the stem, and from here outwards to the point
where they approach the outer hard, woody layer, and that
as they proceed downward in their course in that layer,
they gradually become denser, and of a firm ligneous con-
sistence. When the vascular bundles have reached the
external part of this layer, and become situated beneath
the rind, their thickness is diminished, but not their
firmness and hardness ; the latter peculiarity, however, is
less remarkable, on account of their smaller diameter.
They run in this manner, in the form of slender fibres,
between the firm, woody layer and the rind, to the base
of the stem, or terminate, after a course of variable length,
in other vascular bundles, becoming blended with them.
As all the vascular bundles have a similar course, and the
portion running in the middle of the stem is soft and
herbaceous in all, the medulla-like softness of the centre
of the stems is easily explained. It is also clear that the
hardness of the outer layer of the stem results from the
thickness and solidity acquired by the collective vascular
bundles during tkeir course through this outer layer ;
further, that the liber-like fibrous layer under the rind is
formed by the lower extremities of the vascular bundles,
and is not to be compared with the liber of Dicotyledons.

THE PALM-STEM. 9

The vascular bundles of the Cocos- and Calamus-like
stems are distinguished by their not exhibiting that her-
baceous softness in their course from the leaf to the
centre, and from this to the outer layer of the stem, for
they also appear thick and woody here, although in a less
degree than in the outer layers. With regard to the
inferior portion of the vascular bundle, two varieties are
met with in the Cocos-like stem : it either passes, as in
Kunthia, into a thin fibre, and the external fibrous layer of
the stem is then thin, as in the rest of the forms, or the
vascular bundle divides, at its exit from the hard layer,
into several smaller bundles, which, after a short course,
lose themselves in a multitude of fine fibres ; the fibrous
layer is then thick, as, for instance, in Cocos nucifera,
coronata, &c.

From this course of the vascular bundle is deduced
the following statement : The doctrine laid doivn by
DESFONTAINES, that the new vascular bundles originate in
the centre of the stem, and that the harder and thicker
vascular bundles., situated at the periphery of the stem, are
older than the soft ones occupying the centre, and that,
therefore, the vegetation of Monocotyledons is wholly
different from that of Dicotyledons, is altogether incorrect
and inadmissible.

Obs. 1. From the circumstance that the vascular bundles run from the
leaves to the middle of the stem in a curve of small radius, but that from
here, in their way downward, they only approach the rind gradually, we can
understand how phytotomists have been led to assume that they originate in
the middle of the stem. Indeed, this outward course is not easily observed
in a stem split longitudinally, unless the single vascular bundle is dissected
out. One circumstance, however, must have long since indicated the incor-
rectness of Desfoutaines' doctrine. If, namely, the vascular bundles of the
younger leaves lay more internally in the stem than those which go to the
older leaves, the former could never cross the latter. Now, it is easily
seen in all Palms, that the vascular bundles entering a leaf, cross those which
run to the leaves situated higher up, which is only possible by the arrange-
ment of the fibres described above. This crossing is the more striking the
thicker the stem and the closer its leaves, therefore much more evident in
the species of Cocos than in Kunthia ; it is still more distinct in Xanthorrhoea
hastilis (vide De Candolle, Organogr. tab. 7, 8), in a transverse section of

10 THE PALM-STEM.

the stem of which the vascular bundles entering the leaves have the aspect
of medullary rays. The crossing is also very evident in the stem viPandanus,
Dracaena Draco, Aletris fragrans, Aloe, Bambusa, &c.

Obs. 2. The smaller diameter of the lower liber-like end of the vascular
bundles simply explains the less degree of thickness of the fibrous layer of
the stem. Where each vascular bundle ends in a single filament, as in
Bactris, Geonoma, Lepidocaryum, Calamus, Kunthia, (Enocarpus, Hyospathe,
Rhapis, &c., this layer is very thin ; when, on the other hand, the vascular
bundle gives off several fibres, or when, as in Mauritia vinifera, the fibres
retain a considerable thickness, the thickness of the fibrous layer is not
altogether inconsiderable. I found it in Leopoldinia pulchra from ^ to 2
lines ; in Syagrus cocoides 1 line ; in Cocos nucifera, Euterpe edulis, Mauritia
mnifera, 6 lines thick.

The Cellular Tissue of the Palm-stem.

The cellular tissue is not, as in Dicotyledons, distri-
buted into distinct bark, pith, and medullary rays, since
the vascular bundles are scattered throughout the whole
substance of the stem. Nevertheless, the cellular tissue
exhibits different forms in the different layers of the stem,
which may in many respects be compared with the forms
of the cells of the bark, pith, and medullary rays.

The form of the cellular tissue in Palm-stems in general
possesses but one definite character, namely, it is paren-
chymatous, and its cells are usually arranged in perpen-
dicular rows, the forms of these cells varying much, not
only in different species, but in different layers of the
same stem. In general, these cells are only of a medium
size, and apparently in all species, at a certain period of
vegetation, densely filled with starch.

In the fibrous layer, the cellular tissue is composed of
small, thin -walled cells, mostly expanded transversely,
between which are small intercellular passages. In young
stems, the cortical layer of which is still in full vegetation,
chlorophyll-granules are found in the outer cells, starch-
granules in those lying deeper ; the granular formations
subsequently disappear. The cells of this layer only

THE PALM -STEM. 11

form a perfectly regular tissue where the fibrous bundles
lie far apart ; in most cases the regularity of their arrange-
ment is destroyed by the cells lying next the vascular
bundles having their broad side, or more rarely their
narrow side, directed towards the vascular bundle, in
which latter case a stellate figure is formed round each
bundle (Leopoldinia pulchra).

In that layer of the stem in which lie the thick, hard
vascular bundles, the cellular tissue becomes compressed
into thin lamellae, from the vascular bundles lying very
densely crowded here, and by their mutual pressure fre-
quently (especially in cylindrical stems) forcing them to
assume an angular figure ; these lamellae indeed have a
very variable direction according to the form of the vas-
cular bundle, but taken as a whole, run from without
inward, since the vascular bundles mostly exhibit a form
compressed on both sides. The cells are also here trans-
versely expanded in the direction of the lateral faces of
the vascular bundles, and so much the more, the nearer
they lie to the vascular bundles ; therefore they have in
the cylindrical stems, in which at most only 1 to 3 rows of
cells lie between every two vascular bundles, a very much
elongated form, while in other stems this only occurs
from accidental approximation of vascular bundles, and
the dodecahedral form of the cells reappears in all places
where the vascular bundles lie further apart. In propor-
tion as the cells assume a transversely extended form, the
superposition in perpendicular rows becomes converted
into an arrangement in horizontal rows, so that the cel-
lular tissue assumes the so-called muriform character.
In many Palms, e. g. Cocos botryopliora, the outer vascular
bundles stand behind one another in radiating rows, so
that broad strips of cellular tissue penetrate from 1 to 3
lines deep into the stem in the form of medullary rays.
In these strips, the cells exhibit lateral expansion parallel
to the surface of the stem.

The cells of this layer almost always have much thicker
and harder walls than those of the fibrous laver and the

12 THE PALM-STEM.

interior. The thickening of their walls is usually indeed,
although always noticeable, not very striking, but in cer-
tain Palms, on the contrary, e. g. Diplothemum caudescens,
Cocos botryophora, they attain such thickness as we are
only accustomed to see in wood- and liber-cells. In conse-
quence of this thickening of the walls, the dots (which in
general occur in all Palm-stems) are converted into dis-
tinct canals, which correspond in contiguous cells. In
these two conditions, the thickening of the walls and the
punctation, these cells approach no less than in their
form, the cells of the medullary rays of Dicotyledonous
trees, since these also are constantly thick-walled and
punctated.

The cellular tissue of the central part of the stem ex-
hibits in like manner many variations, which in great
part are connected with the position of the vascular
bundles. In all Palms it agrees in these characters the
cells are thin- walled, in most cases arranged in perpendi-
cular rows ; the cells lying upon the vascular bundles are
mostly somewhat elongated, and depend for the direction
of their transverse diameters on the position of the vas-
cular bundles.

In the interior of the Cocos-\ike stems, the cellular
tissue exhibits a regular parenchyma, the cells are thin-
walled, finely punctated, and only in the investment of
the vascular bundles, or where two bundles lie near
together, do they form transitions to the muriform cellular
tissue, without, however, thereby acquiring thicker walls ;
in Cocos botryopJtora . and Diplothemium caudescens, the
cellular tissue even becomes thinner-walled the nearer it
lies to the centre.

In most Palm-stems, on the other hand, in which the
vascular bundles stand much further apart in the middle
than at the circumference, the cellular tissue of the centre
exhibits considerable differences from that of the outer
layers, becoming very lax, and this in two ways.

In some cases the cellular tissue in the middle of the
stem has very large cells, and thus forms a very soft,

THE PALM-STEM. 13

spongy mass ; e. g. in Geonema simplicifrons, (Enocarpus
minor, Kuntkia montana. In these instances only the
smaller cells, forming the boundaries of the vascular
bundles, retain the form of regular parenchymatous cells ;
the rest, very much enlarged, run out in a radiating
direction from the vascular bundles, and form as many
stellate rosettes as there are vascular bundles. In other
cases, the cells in the central portion of the stem do not
attain to such considerable dimensions ; but the tissue
is rendered lax by the intercellular passages becoming
enlarged into regular air-canals. Calamus forms the
transition here, in which large intercellular passages occur
between the cells in the middle of the stem ; these, how-
ever, still preserve too much of the form of regular
parenchymatous cells, and the intercellular passages are
still too small, to allow of our properly reckoning this
cellular tissue under the so-called compound form. This
occurs to a great extent, however, in Astrocaryum gyna-
canthmn, vulgare, Mauritia vinifera, and especially in
Mauritia armata. Here the cells leave between them
large roundish canals, which run in unbroken continuity
through long tracts in the stem, so that one can blow
smoke through pieces of the stem more than a foot long.
At the extremities these canals are gradually attenuated
till they become completely closed, for the septa of
stellate cells, such as are found in many aquatic plants,
in Musa, &c., do not occur in the Palrns.

Obs. 1. Certain German phytotomists (Heyne, Meyen) have recently
sought to distribute the cellular tissue into a great number of subdivisions^
according to the form of the cells ; this appears to me contrary to nature, on
account of the abundance of intermediate conditions between all these forms.
The above description of the cellular tissue of the Palm-stem may serve as a
testimony that the form of the cell stands in no close connexion with its
function, and that it depends quite as much upon the form, organization,
and position of the adjacent cells and vascular bundles, as on the special
nature of the cells. In comparing the stems of different Palms it is un-
mistakeable that cells, lying in corresponding places in different species, which
have a similar import in the economy of the plants, exhibit wholly different,
often variable forms, and it therefore appears quite improper to attach so
much importance to the form of the cells.

14 THE PALM-STEM.

The Palms are the more fitted to afford this evidence, since the plants not
only form one of the most natural families, but also manifest a very great
similarity, both in respect to their vegetation and their products. The same
conditions, deviation of form of the cells in nearly-allied plants, may also be
demonstrated in other no less natural families, for instance, in the Eerns.

2. I did not find raphides or other crystals in the cells of any Palm-stem.

The Palms have not a bark distinctly separate from the
subjacent parenchyma, and exhibiting a special growth,
such as occurs in Dicotelydonous trees ; but the outer
layers of the cellular tissue are remarkable, and therefore
deserve description. In the young condition they have
thin walls, and cannot be distinguished from the cells of
the subjacent fibrous layer ; in more advanced age, how-
ever, their walls are thicker, and become hard and brown.
In many species for example, in Calamus, in many
species of Geonema this layer remains very thin ; its cells
do not acquire such thick walls, and appear to retain their
vitality throughout the whole life of the plant. In other
species, on the contrary, the surface of which is subject to
destruction by atmospheric influence, as in Cocos, Elais,
the rind acquires a considerable thickness, and gradually
draws a portion of the fibrous layer into its circle. In
such case this is not of equal thickness at all points of
the circumference of the stem, but passes deeply into the
fibrous layer in particular places, while in other situa-
tions it is rather thin. Under these circumstances, the
inner layers of the rind inclose a portion of the fibrous
bundles, which is not usually the case.

The epidermis exists in old age only in the cane-like
and calamoid stems ; in the rest, it is more or less de-
stroyed by the action of the weather. It consists of a
simple layer of minute cells. As a general rule, no sto-
mates occur in it, but they do exist scattered in Ehapis
flabelliformis. In Calamus, it consists of minute cells
elongated in the direction from without inward, and
forms a stony, brittle, shining layer.

The different kinds of pubescence also come under
consideration as appendages of the rind and true cellular

THE PALM-STEM. 15

parts. The youngest portion of the stem is frequently
clothed with a hair-like covering, so long as it is still
young and inclosed by the leaf-sheaths. Sometimes this
appears in the form of actual hairs, which are mostly
closely crowded and coherent into a dense felt ; e. g. in
Bactris tomentosa. In other cases, the covering is com-
posed of scales (ramentd), which exactly resemble those
of the Ferns ; e. g. in Rhapis flabeUiformis, Phoenix
dactylifera. In other instances the cells are combined
into spines of various dimensions. On the leaf-sheaths
and spathes many transitional forms are met with, from
simple hairs, stiff bristles, to strong, hard spines. Spines
of this kind exist on the stems of many Palms, lying
closely appressed to it, so long as the in tern odes are in-
cluded in the leaf-sheaths ; but erecting themselves after
the fall of the leaf, they form a terrible defence to the stem
by their hardness, length, and prickly points.

The spines are sometimes blunt cones, but about an
inch long, as in Mauritia armata ; in Acrocomia sclero-
carpa, Astrocaryum Murumuru, Ayri, gynacanthum, &c.,
on the contrary, they form long, slender, very hard and
acute needles. These are only cellular structures; the
cells of the outer layers are elongated, hard, and have
very thick walls, those in the middle are thin-walled
and parenchymatous ; the middle of the spine is often
hollow.

Structure of the Vascular Bundle.

Before I describe the modifications which the structure
of the vascular bundle undergoes in the different parts of
its course, it may not be out of place to state its com-
position in those situations at which, from the hard peri-
pherical cylinder of wood, it enters, on its way to the
centre, the soft middle substance of the stem. It here
consists of three constituents, which may be clearly dis-
tinguished from each other: 1, of liber; 2, of a bundle
of proper vessels (vasa proprid) ; and 3, of the ligneous

16 THE PALM-STEM.

body. These three constituents are constantly applied
together in such a way that the liber is directed toward
the periphery, the wood toward the centre of the stem,
and the proper vessels lie between the liber and wood.
As a general rule, the following is the minute structure
of the different portions.

The liber (Bast) consists of thick- walled, prosenchy-
matous cells. The opinion of Kieser, therefore (Phyto-
tomie, 209), that the liber-cells of the Monocotyledons
have horizontal septa, was altogether unfounded; and
indeed Moldenhawer (Beitrage, 48) had already found a
prosenchymatous liber in the Grasses. The liber-cells
manifest no definite arrangement in their corresponding
positions ; those lying toward the interior of the vascular
bundle are of the smallest diameter. They have fine
dot-canals (Tiipfelcanalen, canals of the pores).

The proper vessels are composed of a combination of
much elongated cells, with horizontal septa, thin walls,
and of various diameter. The narrower lie partly between
the angles of the wider, partly between the lateral walls
of the others.

The wood consists of elongated parenchymatous cells,
with thin walls, and porous, among which, on the side
directed toward the periphery of the stem, usually lie
two large reticulated vessels, and behind these a variable
number of narrower spiral and annular vessels.

The structure described is not retained unaltered by
the vascular bundle throughout its whole length, but its
structure changes in the different parts of its course in an
analogous manner in all Palms. To demonstrate this,
two methods may be adopted, to investigate either the
transverse section of an entire stem, or an isolated vascular
bundle in the different parts of its course.

In the investigation of the transverse section of a Palm-
stem, we can indeed become acquainted with the struc-
ture of any one of its vascular bundles only at one point
of its course; since, however, each of these vascular
bundles has a definite course from the periphery to the

THE PALM-STEM. 17

centre, and from this again outwards to the periphery,
it is clear that in every transverse section of the stem we
must meet with the fibrous inferior extremities of the
vascular bundles at the periphery ; further in, the thick
and hard part of the bundles formed further up ; and
toward the middle of the stem, the soft part of the
bundles in the situation of their more vertical course
beneath their point of curvature ; finally, we may meet
with the portion of the bundles, in which they run from
the centre toward the leaves, in the most varied situations,
among the others. The last vascular bundles will be cut
through more or less obliquely, the rest in pretty nearly
a fair transverse section.

The examination of such a transverse section by the
microscope, shows that the outer fibrous bundles are
composed solely of thick-walled prosenchymatous cells,
which correspond to the liber-cells of the other vascular
bundles. More toward the interior, we meet with large
bundles, which already exhibit the perfect composition of
the vascular bundle ; they are distinguished by their liber-
mass being relatively very large, and by having the wood
consisting of a single vessel surrounded by but few cells.
The proper vessels likewise exist only in small numbers.
Further inward, where the vascular bundles have attained
their most considerable size, they are composed in greatest
part of thick-walled, lighter or darker brown cells, the
wood is still but slightly developed, yet contains already
one or two vessels of tolerable size, which are inclosed
by few, somewhat thick -walled, cells ; the proper vessels
are also but little developed, and are readily distinguished
from the wood-cells by their thinner walls. Still further
inward, in the transition to the soft part of the stem, the size
of the vascular bundles diminishes ; they exhibit a rounder
form, since the liber-mass is considerably smaller and
assumes the form of a crescent, in the concavity of which
the proper vessels are received, and behind these lies the
strongly developed woody portion. In this occur one or
two large vessels, with several smaller behind them. The

18 THE PALM-STEM.

nearer to the centre the vascular bundle lies, the more
the liber is diminished in mass, till at last it displays only
a very thin crescent, while the size of the woody mass
increases in the inverse ratio, the smaller vessels lying at
its inner side increasing in number. The bundle of
proper vessels enlarges in similar proportion to those of
the woody mass. The softness of the whole vascular
bundle increases with the diminution of the mass of the
liber, because the liber alone contains the thick-walled
elementary organs.

Similar changes in the structure of the vascular bundle
are met with, when it is dissected out from the stem and
examined in different parts. In this way we may not
only obtain, by comparison of transverse sections of one
and the same vascular bundle, a survey of the changes of
its size and structure which leaves no room for doubt,
but we may detect more readily than in the cross section
of an entire stem, the changes which the vascular bundle
undergoes in its way from the centre of the stem to the
base of the leaf.*

These changes are as follows : the nearer the vascular
bundle approaches to the leaf, the more the liber-mass
diminishes in size and the woody portion increases, a
great multiplication of the vessels of the latter being con-
nected with this, these, however, considerably decrease
in size. In the vicinity of the point of emergence from
the stem, a division of the vascular bundle into several
(up to six) portions already begins to be effected, this
taking place in such a manner, that small bundles of liber
appear on the outer borders of the woody portion, at its
posterior and lateral surfaces, behind which, and at some
distance higher up, are found the rest of the systems
(wood and proper vessels) belonging to a perfect vascular
bundle, so that the entire vascular bundle consists of a
circle of smaller bundles, which all have their woody por-

* What relates to these two methods of investigation is not a translation
of the original text, for the latter referred specially to the illustrations here,
and therefore would be incomprehensible without them. (H. v. Mohl.)

THE PALM-STEM. 19

tions directed toward a common centre, and by the simple
separation of these subdivisions, it becomes decomposed
into just so many bundles containing all the essential parts
of the vascular bundle.

From the character of the vascular bundle, as above
described, follows incontrovertibly the total falsity of the
generally received opinion, that the thicker and firmer
vascular bundles lying in the outer parts of the stem are
the older and lignified, and that the softer, lying in the
middle, are the younger, not yet come to their complete
development.

Modifications of the Structure of the Vascular Bundle in
various Palm-stems*

Although the structure of the vascular bundle exhibits
a common type in all Palms, modifications occur in the
different species.

In the cane-like stems, the liber of the outer hard
layer of the stem displays a very considerable develop-
ment ; toward the centre it, indeed, diminishes again in
volume, yet still retains a moderate size, and since the
the vascular bundles are not very distinct from each
other, the middle of the stem possesses tolerable solidity.

In the cylindrical stems, the liber of the outer layers ex-
hibits the greatest development that occurs in the Palms,
into a mass often elongated in the direction from within
outwards ; in the middle of the stem the vascular bundle
acquires an herbaceous softness, partly through the dimi-
nution of the liber-bundle, partly through the walls of
its cells becoming so very thin, that in a cross section
they resemble parenchyma-cells.

In the cocos-like stems the liber only increases slowly
in the passage of the vascular bundle from the fibrous
layer toward the interior, and does not attain any con-
siderable size ; the vascular bundles of the outer parts of

* This portion consists of extracts, since the detail in the original referred
to the illustrations. (H. v. Mohl.)

20 THE PALM-STEM.

the stem are also in a less crowded condition than in the
two preceding forms. Partly by this, and partly by the
smaller amount of decrease of the liber in the middle of
the stem, is explained the more uniform hardness of the
different layers of the latter.

The vascular bundles of Calamus exhibit a very pecu-
liar structure. The liber is here also strongly developed
in the outer layer of the stem, but the woody portion
displays this peculiarity, that, disregarding rare excep-
tions, instead of several large vessels, it contains only
one of unusual dimensions, occupying the centre of the
bundle. Behind this large vessel (except in the outer-
most vascular bundles) lie small spiral vessels. The cells
of the woody portion have thick walls, and thus may
readily be confounded with the liber-cells in a cross sec-
tion. The proper vessels are distributed in two groups,
which, with the spiral vessels, form as it were the points
of a triangle inclosing the large vessel.

Of the Structure of the particular Anatomical Systems
of the Vascular Bundle.

The cells of that part which I denominate liber always
have a diagonal septum. In a young condition, they,
like all other thick- walled cells, are composed of delicate,
colourless membrane. When with increased age they
have become thicker, they afford clear evidence that the
membrane of the vegetable cell grows in thickness by the
deposition of layers. In transverse sections of the walls
of the liber-cells of all Palms, delicate concentric lines
may be seen, and that these lines form the boundaries
of the different layers composing the cell-membrane, is
manifest from the fact that sometimes, when the section
has been made with a razor which is not very sharp, these
layers separate from each other, and the slice of cell-
membrane appears in the form of distinct concentric
rings. Frequently the colour of these cells is not uniform
throughout the whole thickness of the membrane, and

THE PALM-STEM. 21

some of the component layers are often of a darker hue
than the rest.

A second remarkable peculiarity of these cells is their
porosity. Both in the transverse and longitudinal sec-
tions we see fine striae, which run from the cavity toward
the outer surface of the cell. When a high magnifying
power is used, there remains no doubt that these striae
are canals perforating the cell-wall. As a general rule,
their diameter does not exceed ^~ of a line. As in the
case of the pores of cellular tissue, the canals of adja-
cent cells correspond to each other.

The second constituent of the vascular bundle, which
I have termed wood, is composed of two organic systems,
cellular tissue and vessels.

The cellular tissue of this woody portion consists of
colourless parenchymatous cells, the walls of which are
not very thick. They are usually somewhat elongated,
stand in vertical series one above another, with horizontal
septa ; never lie in series diverging like a fan from the
hindmost point of the woody mass, but form an irregular
parenchyma, the cells of which, in the vicinity of the
vessels, are arranged according to the form and position
of these latter. These cells never contain starch-granules ;
their walls are studded with large and small pores like
the cells of Cycas.

The woody mass, as already mentioned, always lies at
the inner side of the vascular bundle ; but in the transi-
tion of the fibrous bundle, devoid of vessels, into the
condition of vascular bundle, it is very frequent, and
almost the rule, for the woody mass to lie, not at the
inner side, but in the middle of the liber-bundle. In the
vascular bundles of the outer, hard layers of the stem,
also, a narrow strip of liber-tubes often runs round the
posterior face of the woody mass, so that this is completely
surrounded by liber- tubes.

In other cases the membrane of the wood-cells is itself
thickened, and thus they acquire a resemblance, at least
in the cross section, to the liber-cells ; however, they are

22 THE PALM-STEM.

mostly to be distinguished from these by their larger
cavity and somewhat thinner walls, and in the longi-
tudinal section by the septum being, at all events in the
vicinity of the vascular bundle, horizontal. This thicken-
ing occurs sometimes here and there in the outer bundles
of many Palm-stems, e. g. in Runtliia montana, in which
case it is met with in one vascular bundle and not in the
others ; or it is a structure occurring regularly in all
bundles, which, however, is only the case in Calamus.
Notwithstanding that there is here a great similarity
produced to the liber-cells by the thickening of the walls,
they may be distinguished from these by the somewhat
thinner walls, a larger cavity, as well as by the circum-
stance that, with the exception of the hindermost, they
are elongated in a direction parallel to the wall of the
large vessel. Their walls, like those of the liber-tubes,
consist of several layers, and possess pore-canals, which
are particularly striking in longitudinal sections, since
from their small distance from each other, the cell-wall
cut through possesses almost a moniliform aspect ; and
under these circumstances the nature of these canals, as
excavations perforating the cell-wall down to the outer-
most layer, may be recognised most clearly.

The vessels of the Palms must be divided into the
large and small. Each of these kinds, as is clear from
what has been said already, occupies a definite place in
the vascular bundle. The large vessels, traced from the
lower fibrous extremity of the vascular bundle to its exit
into the leaf, do not anywhere exhibit the form of the
spiral, but that of the scalariform or reticulated vessel.
These vessels are composed of rather short tubes, standing
one above another. This composition may be perceived
even by the naked eye in the very wide vessels of Cala-
mus Draco, Mauritia vinifera, &c., the length of one of
the tubes in these plants amounting to 1 2 lines. The
face where the ends of the tubes meet one another is very
seldom horizontal, but mostly inclined considerably to-
ward the axis of the vessel. As a general rule, the ends

THE PALM-STEM. 23

of the tubes do not lie one behind another in the direc-
tion of a line drawn from the periphery to the centre of
the stem, but side by side. The tubes do not always
succeed one another singly in the longitudinal arrange-
ment, for frequently two or three stand next below a
single one, so that in the cross section there appear to be
2 3 vessels side by side. This occurs sometimes even
low down in the vascular bundle, in which case these
vessels often again blend into a single one. Some of
these vessels attain a very considerable size. In the
lower part of the course of the vascular bundle, they
have indeed scarcely a diameter of ^ of a line ; but the
vessels occurring in the middle tract are among the
largest that are met with in the vegetable kingdom :
thus the vessels of Bactris mitis exhibit a diameter of
from ^ to of a line, those of Desmoneus mitis, (Enocar-
pus minor, ~ to ^, of Astrocaryum gynacanthum ~ to ^,
Corypha cerifera and Mauritia armata to ~, Mauritia
vim/era \ to \, Calamus Draco i to J of a line.

The walls of these vessels universally exhibit the form
of a dotted tube.* But the nature of this varies according
to the nature of the adjacent parts. As this circumstance
has almost entirely escaped phytotomists, a more minute
exposition of it may not be out of place here. I have
already pointed out, that in cellular tissue the position of
the pores of one cell exerts a definite influence over the
position of the pores of the adjacent cells, and that the
pores of two contiguous cells always correspond. Now this
occurs also in the reticulated vessels. It is a universal
law in the reticulated vessels and scalariform ducts of all
plants, that the dots and slits of their walls are somewhat
shorter than the breadth of the cell or vessel in contact

* In the Latin original I have called them vasa porosa, v. punctata,
because the term reticulated vessel did not appear to me adapted to the
kind of punctation. But since I have subsequently found that the pores of
dotted vessels of the Dicotyledons are distinguished from those of the vessels
of Palms, by the fact that there is a cavity between each pair of pores in the
former, which is wanting here, I now select the expression reticulated vessel,
to avoid the necessity of making a new term.

24 THE PALM-STEM.

with them. Since, as a rule, the vessels are surrounded
by elongated cells, and the lateral walls of these cells,
perpendicular to the vessel, run down some distance ver-
tically upon it, this causes the well-known appearance
that the dots or slits lie in a straight row, one above
another, in the vessel. In the Palms, the vessels are, in
most cases, surrounded by short, dodecahedral paren-
chyma; since, therefore, in accordance with the above
law, the pores are only formed at those places in the
vessel at which the parallel walls of the neighbouring
cells are grown to the wall of the vessels, these vessels
exhibit an apparently irregularly grouped distribution of
the pores and, between these groups of pores, free inter-
spaces, which correspond to the perpendicularly-placed
side and cross walls of the adjacent cells.

In other cases, where elongated cells are in contact with
the vessel, the pores lie in regular vertical rows. When
two vessels are directly applied together, the pores of the
lateral walls, grown to the other vessel, assume the form
of transverse slits, which are as long as the breadth of
this wall, whereby the vessel becomes scalariform, while
the other sides, in contact with cells, possess the form of
the reticulated tube. These relative conditions are not
peculiar to the Palms, but occur in the same manner in
other plants, for instance, most distinctly in the Tree-
ferns.

Obs. It is further to be noticed, that cases also occur in which the pores
have not the whole breadth of the adjacent elementary organ. For instance,
it not unfrequently happens that the pores are a good deal shorter, and lie in
regular horizontal lines, intermingled with longer slits. I found a peculiar
deviation from the general rule, that the pores of two adjacent vessels
correspond exactly in position, form, and size, in Corypha cerifera, where
one vessel was studded with longer slits, while the other possessed rows of
roundish or elliptical pores corresponding to these slits.

Both the investigation of full-grown vessels and their
development, presently to be discussed, prove that the
pores and slits are not actual openings, but are closed
by a delicate membrane at the outside. This may be

THE PALM-STEM. 25

perceived most clearly when a longitudinal section passes
through the adjacent walls of two scalariform ducts, in
which case we recognise distinctly that the slits are actual
excavations in the walls of the vessels, and not elevations,
as stated by Bernhardi (Ueber Pflanzengefasse, p. 35),
Treviranus (Beitrage, p. 22), and Meyen (Phytotomie,
p. 253), but that the interspaces between the slits form
elevations which project into the vessel. It may likewise
readily be perceived that a membrane is stretched between
the fibres, separating the fibres of the two vessels under
the from of a simple, dark line. The development of
these vessels shows that this membrane forms a tube
inclosing the fibres. The pores and slits equally exhibit
an evident rim, which does not depend, as Mirbel assumes,
on the presence of a projecting border, but is, on the con-
trary, caused by the borders of the slit being truncated
by an oblique surface.

In many cases, as in Calamus for example, the place
where two vascular tubes are connected is conspicuous,
from each tube ending in a broad ring ; the two adjacent
rings form a band surrounding the vessel, as Moldenhawer
has shown long since in other plants. When the tubes
meet by such a ring, they are freely open to each other.
In other cases, on the contrary, particularly in Desmoneus
mitis, Cocos nucifera, Mauritia vinifera, armata, Kunthia
montana, Astrocaryum gynacanthum, vulgar e y Coryplia fri-
gida, such rings are seen at the points of connexion of the
tubes, and in these are found septa. The existence of
such septa is indeed denied utterly by all phytotomists ;
but having observed them hundreds of times, not only in
the Palms, but in many other Monocotyledons, and even
in many cases in the porous vessels of Dicotyledons, I do
not hesitate to assert their existence in the most positive
manner. The direction of these septa is usually such
that we obtain a front view of them in a longitudinal
section made in the direction of the radius of the stem.
They differ completely from all other membranes of plants,
in being formed of a reticulation of thick fibres, with
large openings between them. In the stems of Palms

26 THE PALM-STEM.

where these septa cut the axis of the vessel obliquely, and
therefore are elliptical in form, the fibres are mostly hori-
zontal. At the two rounded extremities of the septum
the orifices present the form of little slits or round holes,
in the middle of broad slits or oval openings, and at both
sides of the septum likewise occur smaller, roundish, or
ovate orifices. Each of these orifices is bounded by a
narrow rim. In other cases, the orifices exhibit the form
of narrow, transverse slits, giving the septum quite a
scalariform aspect. The openings are usually actual per-
forations, very rarely a thin membrane is stretched over
them. The fibres of the septa are double, and the rim
originates from the same cause as in the scalariform
ducts. These septa occur very frequently in the Palms ; in
many species, however, as for instance in Cocos nucifera,
they do not occur at every point of junction of two vessels,
for some of these end in the ring above described. To
avoid the necessity of recurring to these septa in the
examination of the root of the Palms, I will mention here,
at once, that in the large vessels of the roots the septa
are usually perpendicular to the axis of the vessel, and
therefore of roundish shape. In this case they are not
scalariform, but reticulated, perforated by large, roundish,
and many small punctiform orifices. The course of the
fibres does not always correspond exactly in the two com-
ponent plates of the septum, whence a portion of one
plate often projects into the opening of the other.

In the porous vessels of the Dicotyledons it is well
known that vesicular cells often occur, which Kieser be-
lieved to be composed of the same membrane which forms
the wall of the porous vessel, whence he assumed (Phy-
totomie, p. 237,) that such vesicles could by no means
occur in the Monocotyledons. I however found, though
indeed but seldom, vesicular cells, similar to those of the
Dicotyledons, in the large vessels of the Palms, for in-
stance, in Corypha cerifera*

* I have not traced the development of these cells in the Palms. Doubt-
less they have the same character as in the Dicotyledons, in regard to which
from recent researches, I think that I am not wrong in assuming that they

THE PALM-STEM. 27

The course of development of these vessels, which I
investigated in the germinating date-palm, in the apex of
the stem of Rhapis flabelliformis, and in the root of
many species of Palms, and with which the development
of the large vessels of Dioscorea, Tamus elephantipes, &c.,
fully agrees, is as follows : In young shoots, we find in
the situation where the large vessels subsequently lie,
perfectly closed, large, and cylindrical tubes, which are
composed of a colourless and very delicate membrane. In
tubes a little older, we find a network of very delicate,
transparent fibres upon the internal surface ; these have
a horizontal direction, are connected at places which cor-
respond to the longitudinal septa of the adjacent cells by
perpendicular and oblique fibres. As a general rule, the
horizontal fibres are so arranged that they are not con-
tinuous over several lateral walls of the vessel, but termi-
nate where they reach a longitudinal septum of an adjacent
cell, and here pass into a fibre, going obliquely upward
or downward, so that a mesh of a fibrous network comes
as a direct successor of the fibre on the adjoining side-
wall of the vessel. From this it is most clear that these
vessels are not originally spiral vessels, the fibres of which
become reticularly connected by the subsequent develop-
ment of cross-fibres. This is made still more evident by
the fact, that in the first origin of the fibrous network,
we in many cases meet with the fibres only perfected at
particular places, the walls of the vessel appearing still
perfectly smooth in other situations. The older the vessel
grows, the broader and thicker its fibres become, and
the interspaces between them are diminished in width in
proportion, till at last they display themselves as mere
narrow slits. The septa are perfected in a manner wholly
analogous, but the original delicate membrane is generally
destroyed, after a time, in the interspaces of the fibrous
network.

are produced by a protruding expansion (a kind of hernia) of the adjacent
cell, which penetrates the pore, and either tears through or causes the
absorption of the primary membrane of the vessel.

28 THE PALM-STEM.

That these vessels, though not the result of a meta-
morphosis of spiral vessels, belong to one and the same
system, is evident from the fact, that spiral vessels occur
in many Monocotyledons in the situation where the reti-
cular vessels lie in the Palms, and in the Grasses, as
Moldenhawer showed, the same row of tubes is developed
at certain places into spiral, and at others into porous
vessels.

The smaller vessels lying behind the reticulated are never
reticulated, but always spiral or annular vessels, be the
stem under examination as old as it may. The number
of annular vessels in each vascular bundle is but small,
in general only spiral vessels exist. When annular ves-
sels do occur, they are always situated the farthest back in
the bundle, the vessels lying nearer to the large vessels
being always spiral. The turns of the spiral are always
far apart, especially in those vessels which are farthest from
the large vessels. It is easy to satisfy one's self of the
presence of an outer membrane inclosing the spiral fibre.

I have mentioned above, under the name of proper
vessels (vasa propria), a bundle of thin-walled cells lying
between the wood and liber, as the third constituent of
the vascular bundle. This bundle is distinguishable from
the surrounding cells, both by the thinner walls of its
elementary organs, and by narrow and wide cells lying
intermingled in it. This part opposes considerable diffi-
culty to anatomical investigation, on account of its soft
texture, and also its great transparency. This circum-
stance explains why these proper vessels, although they
occur in most of the Monocotyledons and in a portion of
the Dicotyledons, have been overlooked by almost all
phytotomists, for Moldenhawer is almost the only one
who was accurately acquainted with them, in Zea Mays
and Bambusa, while Amici saw them in Calamus, but did
not recognise their nature, and Kieser, who likewise saw
them in Calamus, explained them as spiral vessels ;
Bernhardi and Meyen, the latter of whom found these
proper vessels in Scirpus lacuslris and some other Mono-

THE PALM-STEM. 29

cotyledons, regarded them as prosenchymatous cells, and
distinguished them neither from the liber nor the wood.
In a longitudinal section we perceive that the cells are
elongated in these bundles, and stand one above another
mostly with horizontal septa ; the septa are, however,
sometimes oblique, and the tubes not unfrequently ter-
minate in a point. We may distinguish narrow and wide
cells, frequently with a tolerably strong difference of
diameter. Moldenhawer (Beitrage, p. 126) regarded the
wider tubes as cells of the usual kind, and the narrow as
proper vessels, which he referred to the same system as
the milk-vessels of Chelidonium, Asclepias, &c. So far
as my own investigations showed, I cannot agree with
Moldenhawer in this distinction of wide and narrow
tubes, for I frequently believed that I saw both contain-
ing an opaque, thickish sap, in which swam a great
abundance of fine granules ; I am, therefore, compelled
to refer both to the same system. I never found the
contained sap milk-white, but only more or less opalescent.
I never could observe any currents, but only an oscilla-
tion of the minute granules, which appeared to me merely
molecular motion. The septa between the tubes are
completely closed. In the stem, the wide and narrow
tubes are intermingled without order in the entire bundle,
but a different condition will be met with in the root.
In Calamus, the bundle of these proper vessels is not
only divided into two separate portions, but the narrower
tubes are frequently separated from the wider, as the
latter often lie isolated, at the limit between the wood-
and liber-cells.

I have denominated these tubes proper vessels, because
the nature of their contents brings them nearest to the
system known as the vital (Lebensgefasse) or milk-ves-
sels ; I shall bring proofs further on that they are different
from this system. In reference to their development,
I can merely state that, in the Palms as in other Mono-
cotyledons, they precede the formation of the woody por-
tion in so far that, when a fibrous bundle devoid of vessels

30 THE PALM-STEM.

is traced upward to the point where it displays itself as a
perfect vascular bundle, they show themselves sooner than
the woody mass itself. However, this earlier develop-
ment relates only to space and not to time. I do not
venture to give an opinion as to the function of this sys-
tem, and for the present am content to have demon-
strated their existence anatomically.

Comparison of the Palm-stem with the stems of other
Monocotyledons.

With regard to the course of the vascular bundle, the
Palms stand nearest to Dracana, Aletris, and Aloe ; with
reference to the organization of the bundle, to the Grasses.

As to the structure of the vascular bundle of the
Grasses, I refer to the excellent researches of Moldenhawer,
and merely mention here, that in the Grasses the vascular
bundle not only increases in size from the periphery to-
ward the centre of the stem, but its structure is altered
at the same time, in a manner analogous to what occurs
in the Palms. Beneath the surface of the stem we meet
with fibrous bundles without vessels, then fibrous bundles
containing a bundle of proper vessels, and these are
adjoined by vascular bundles with reticulated vessels,
while small vessels occur, in addition, in the inner. The
perfect vascular bundles, for example, of Arundo Donaoc,
contain two large vessels surrounded by thin-walled wood-
cells, behind these lie a row of two or three small vessels ;
the hindmost portion of the wood-cells is composed of a
mass of thick-walled, dotted prosenchyma, as also fre-
quently occurs in the leaves and other organs of the
Palms. Between the wood and the liber lies a bundle of
proper vessels, the large cells of which are mostly qua-
drangular, and have a narrower cell lying between their
angles.

In Dractena and Aloe, the former of which is especially
interesting on account of the minute researches on its

THE PALM-STEM. 31

growth in thickness by Dupetit-Thouars, we meet with a
structure of the vascular bundle which is aberrant in
many respects. In a cross section of the stem of these
plants, two evidently distinct layers may be detected, the
inner being soft and pith-like, and acquiring no increase
of size with the age of the plant, while the outer forms a
firm mass, and gradually increases in thickness with the
age of the plant. Anatomical investigation demonstrates
that the inner soft substance has wholly the structure of
the Palm-stem, its vascular bundles running in the same
way from the periphery to the centre, and from this to
the leaf. The most external of these bundles are smaller,
poor in vessels, and closely crowded, the inner consist of
several rows of dotted liber-tubes, a bundle of proper
vessels, and a woody portion, which differs from that of
the Palms only in so far that its large vessels are not iso-
lated, but united with the smaller vessels into the form of
a crescent. Tracing these vascular bundles downwards,
we see them enter the outer firm layer of the stem ;
instead, however, of running down the stem in the form
of fine fibres into a laxer cellular tissue, as in the Palms,
they retain a considerable diameter, notwithstanding that
they lose their woody portion, since the prosenchymatous
cells of which they are composed are very wide. They
contain a small bundle of proper vessels in the centre.
Under this form they run downward in the stem, as in
the Palms, but are not isolated, for they unite together
laterally into a network, like the ligneous bundles of the
Dicotyledons. The parenchyma in which they are im-
bedded consists of tolerably thick-walled, porous cells,
elongated in the radial direction. This outer firm layer
therefore corresponds to the fibrous layer of the Palms.
No trace of it occurs at the apex of the stem, since it is
composed of the lower extremities of vascular bundles*:
the lower down in the stem we examine, the thicker
this layer appears, and thus the stem is conical and not
cylindrical. The younger layers of the evascular fibrous
bundles are applied upon the outer side of the older, and

32 THE PALM-STEM.

hence injured places upon the stem, over which no new
layers are deposited, are gradually converted into depres-
sions. On account of this continuous deposition of new
layers, the assertion made by Dupetit-Thouars (premier
Essai) and received into all botanical works, that the stem
of Draccena does not increase in thickness so long as it
remains without branches, is incorrect, and it is easy to
convince one's self of this by the comparison of young
and old specimens ; thus, I found that the stem of a
Dracaena Draco, two feet high, was only about one inch
thick, while trunks twenty to thirty feet high had attained
a thickness of from four to six feet, although devoid of
branches.

Aloe has exactly the same structure as Draccena, only
the outer firm layer is very thin in proportion to the
soft centre. Thus, in a stem of Aloe glauca two inches
thick, it was only two lines ; while in a stem of Dractena
Draco, two inches thick, it had already reached a thick-
ness of six lines. The vascular bundles of Aloe resemble
those of the Palms still less, for the vessels are scattered
irregularly in the woody portion ; however, the anterior
always have the form of reticulated, and the posterior of
spiral vessels.

Comparing the stem of other Monocotyledons with
that of the Palms, in reference to the course and organi-
zation of the vascular bundles, we find in all (excepting
the very lowest forms), if not so great a similarity as in
Drac&na, yet a great analogy. It is, namely, common to
all Monocotyledons for the small evascular bundles to
lie at the periphery of the stem, while those lying further
in contain a bundle of proper vessels, to which succeed
bundles possessing large and small vessels, while the fully
developed lie toward the centre of the stem.

These bundles exhibit more or less deviation from those of the Palms, in
reference to their intimate structure ; but that they are all formed after one
and the same type is unmistakeable. In general, the liber-layer is developed
in a much less degree than in the Palms ; for in many only the outer small
bundles exhibit an investment of thick-walled prosenchymatous cells, while

THE PALM-STEM. 33

the inner possess but a few thin-walled, wide liber-cells, scarcely to be dis-
tinguished from the parenchyma- cells and proper vessels in the transverse
section, e. g. in Asparagus officinalis, Lilium bulbiferum, Orchis militant, and
Sagittaria sagittifolia. When the inner bundles do possess thick-walled
liber-cells, these exist but in small quantity, and form only a narrow crescent.
Sometimes their diameter is so great in proportion to their thin walls, that
these cells are only to be recognised as the organ so greatly developed in the
Palms, by their position and their greater development in the outer bundles ;
for example, in Nusa paradisiaca, Hemerocallis flava, Tulipa gesneriana,
Fritillaria imperialis, Ruscus Hypophyllum, Iris sibirica, Epidendron elonga-
tum, Aloe Commelini, and Scirpus lacustris.

The proper vessels occur throughout the whole series of Monocotyledons,
and lie in the same position between the liber and wood as in the Palms ;
they also possess the same structure, and contain the same opaque sap. In
many plants they are more developed in reference to number and size than in
the Palms ; e. g. in Asparagus officinalis, where single ones, lying toward the
inner side of the bundle, exhibit uncommonly large cavities. In Mma
paradisiaca, the proper vessels, with their thin investment of liber-cells, form
a large bundle, lying separate, in part, from the woody bundle, and not
much inferior to it in size. In Dioscorea villosa also, and Tamus, some of
the vessels lying toward the interior are developed to an unusual size. In
Sagittaria sagittifolia the proper vessels form about half of the entire vas-
cular bundle, and are difficult to discriminate from the surrounding cells,
since these likewise have very thin walls.

The woody portion consists, as in the Palms, of more or less elongated
parenchymatous cells ; its vessels may also be divided into large and small ;
but the large are not so much separated from the small in their position in
most Monocotyledons, for all the vessels generally lie together. A resem-
blance to the vascular bundle of the Palms does, however, occur here in so
far that the large vessels lie in front and at the two sides, and the smaller
behind and between the former, so that the total mass of vessels forms a
more or less regular crescent, with its concavity outward, as for instance in
Asparagus officinalis, Convallaria Polygonatum, and Lilium bulbiferum. In
all these cases the large vessels lying at the two extremities of the crescent
are reticulated tubes, while the smaller, lying further back, have the form
of scalariform ducts, and the smallest, most posterior, that of spiral or an-
nular vessels. Ruscus Hypophyllum makes an exception to this arrange-
ment, as the largest vessels here lie behind and toward the interior. One
consequence of this crescentic position of the vessels is, that the bundle of
proper vessels retreats back between the horns of the crescent, and thus is
half inclosed by the wood ; e. g. in Asparagus, Convallaria Polygonatum, and
Lilium bulbiferum. This is the case in the highest degree in Tamus and
Dioscorea, where the proper vessels are retracted so much between the

34 THE PALM-STEM.

crescent formed by the vessels, that the large vessels come m contact again
in front of the bundle of proper vessels.

The unusual form of these vascular bundles may be an apology for my
entering somewhat more minutely into their structure. The vascular bundles
of these plants lie in two circles alternating with each other, those of the
inner being considerably larger. Each of these vascular bundles is composed
of a combination of three vascular bundles : one, the largest of them, situ-
ated internally, consists of a crescent of vessels, the most anterior and largest
of which are finely dotted reticulated vessels, while only the most posterior
and smallest have the form of spiral vessels. The proper vessels belong to
this bundle. In front of this occur two vessels of tolerable size, which
in many cases are likewise connected by a number of smaller vessels into a
crescent, the convexity of which is directed outwards. Behind these vessels
lies a second bundle of proper vessels. It is therefore evident that each of
these vascular bundles is formed by the blending of two vascular bundles.
The structure of the vascular bundles of the outer circle, and of those formed
in the slenderest shoots of Tamus Mepliantipes, show clearly that the outer
and smaller of these vascular bundles is also composed of two bundles ;
for in the vascular bundles of the outer circle, the crescent formed by the
vessels of the larger bundle, looking inward, is more widely open, and the
anterior vascular bundles separate into two, each of which possesses a bundle
of proper vessels at its inner side. In the young shoots of Tamus Mephan-
tipes, the structure approaches still nearer to that usual in the Monocotyle-
dons, for the smaller bundle is either altogether wanting, or, if present, is
likewise divided into two portions, which, however, do not converge toward
the large vessels, as in the stem, but have their bundle of proper vessels
lying at their inner side.

In the forms enumerated hitherto, the analogy of the vascular bundle
with that of the Palms is so evident as to require no further demonstration
in that respect. The forms in which the different vessels exhibit almost
equal diameter e. g. in Henierocallis flava, Tulipa gesneriana, Fritillaria
imperialist Orchis militaris, Iris sibirica, and Aloe Commelini, are rather more
removed ; but here also the most anterior vessels are constantly reticulated,
the posterior annular and spiral vessels.

It may not be superfluous to mention a structure which may readily give
rise to errors. In the vascular bundles of many aquatic plants an air-passage
occurs, possessing no proper walls ; for example, in Alisma Plantago, Sagit-
taria sagittifolia, Scirpus lacustris, Cyperus Papyrus, &c. If merely a trans-
verse section of these vessels be examined, this canal may easily be taken
for a vessel, as happened with Bernhardi (iiber Pflanzengefasse, p. 16, in
Alisma), and Meyen (Phytot. pi. vi, fig. 1, in Scirpus). The whole form of
the vascular bundle becomes altered through this canal. In Scirpus and
Cyperus this is not so much the case, the vascular bundle being in general

THE PALM-STEM. 35

very similar to that of the Grasses ; but in Sagittaria, in which exist many
vessels of tolerably equal size, these accommodate their position to the air-
canal, and form a crescent, convex to the outer side, enveloped by a large
bundle of proper vessels. This is the case also in Alisma Plantago ; only a
portion of the vessels are also scattered irregularly in the woody portion.
I did not find proper vessels in the vascular bundles of this plant, but the
liber-tubes immediately adjoined the wood.

As I have already observed, the large vessels of the Monocotyledons, as
a general rule, exhibit the form of reticulated vessels ; but that this does
not hold without any exception, is shown by the masterly researches of
Moldenhawer, on the vascular structure of the Grasses. In these, however,
the vessels are still reticulated in the greater portion of their course ; in
others, they exhibit the form of spiral vessels. Thus, in the petiole of
Musa paradisiaca, a very large vessel occurs in the middle of the woody
bundle, in place of which, in the middle vascular bundles of the stem, three
or four vessels occur. Now this vessel, as a general rule, exhibits the form
of a spiral vessel, with many parallel fibres ; and only in rare cases for in-
stance in the lowest parts of the vagina of the leaf and in the rhizome did
I find the fibres of this vessel blended together, and this frequently merely
at particular places. In the same way I found, as a rule, only spiral vessels
in the stems of Ti/pha angustifolia, Sparganium ramosum, and in the petiole
of Calla JSthiopica.

I shall subjoin but a few words on the cellular tissue of the Monocotyle-
donous stem. It consists of large cells, mostly with thin walls, yet fre-
quently having pores ; they have intercellular passages between them, and
exhibit a transition of form between the polyhedral and cylindrical. These
cells become of smaller diameter towards the surface of the stem, with
which is combined an increased thickness of the walls. This thickening
occurs to such an extent in many Monocotyledons, at the place where the
outer vascular bundles lie, that a firm ring is formed around the stem, for
instance, in Arundo Donax, Ruscus Hypophyllum, Asparagus qfficinalis, Con-
vallaria Polygonatum, Lilium bulbiferum, Iris sibirica, Dioscorea villosa,
Tamils Elephantipes, Sparganium ramosum, Triglochin palustre, Alisma
Plantago, &c. From their thickened walls, their smaller diameter, and
elongated form, these cells resemble the liber-cells, but it would be a great
mistake to compare this ring with the liber of the Dicotyledons, since :
1. There are many plants, e. g. Fritillaria imperialis and Tulipa gesneriana,
in which these cells are wide and less thickened in the walls, forming a
distinct transition into the parenchymatous cells. 2. This ring is not
sharply defined at its inner side, but passes gradually into the parenchyma
of the stem. 3. The relation to the vascular bundles and the leaves is
altogether different from that of the liber-bundles of the Dicotyledons to
these. 4. Many herbaceous Dicotyledons exhibit this ring and liber-

36 THE PALM-STEM.

bundles together with it. From these reasons it is inadmissible, although
Link (Grundlehren, p. 143 ; Elem. Phil. Bot. p. 140) explains this ring as
liber, a view which Kieser also propounded (Phytot. p. 72). This ring is
very sharply denned at its outer side, and surrounded by parenchyma which
represents the bark.

I have already expressed rny opinion that the proper vessels lying in the
vascular bundles are not to be reckoned as belonging to the milk-vessel
system, in which the so-called vital sap is contained in Chelidonium, Asclepias,
Euphorbia, Musa, Fic^ts, &c. This is clear from the fact that the vital-sap
vessels do not occur in the place in which these proper vessels lie in the
Monocotyledons, for the former lie isolated in the interspaces of the cellular
tissue, chiefly in the vicinity of the vascular bundles, in the pith and the
bark. But it is best proved by the existence of many Monocotyledons, in
which both these kinds of vessel are met with, independently, each in its
proper place, and in these cases the two kinds of vessel conveying wholly
different sap. Thus it is known, for example, from Moldenhawer's
researches (Beitrage, p. 134), that milk-vessels lie in the parenchyma in the
vicinity of the vascular bundles in Miisa, the sap of which, like that of
Sambucus, assumes a red colour, and which was also recognised as vital sap
by Schultz (Natur. d. leb. Pfl. i, 537). These vessels, then, are distinguished
from the proper vessels in the vascular bundles by their different situation
and by the red colour of their contents, which colour is never assumed by
the sap of the proper vessels. The proper vessels in the vascular bundle of
Sagittaria are no less clearly distinguished from the milk-vessels lying scat-
tered in the parenchyma of the peduncle and petiole, for the latter convey a
milky, the former a very transparent sap. In Alisma Plantago, likewise, the
vessels filled with milky juice are wholly detached from the vascular bundles.

Comparison of the Vascular Bundle of the Palms with
that of the Dicotyledons.

In order to place in a clearer light the connexion of the
organization of Palms with that of the Dicotyledons, and
in order to state more accurately the grounds on which I
have called the different parts of the vascular bundle of the
Palms, liber and wood, a comparison of the vascular
bundle of the Palms with that of the Dicotyledons be-
comes necessary, since in the latter alone can there exist
no doubt as to the true import of its parts.

In herbaceous Dicotyledons the vascular bundles stand

THE PALM -STEM. 37

in a circle, and are separated from each other by strips
of parenchymatous cells varying in breadth ; the vascular
bundle of Laserpitium aquilegifolmm may serve as an
example. We find in this a great quantity of irregu-
larly distributed vessels, the larger of which lie toward
the outside, the smaller toward the interior ; the exterior
vessels are porous and scalariform ducts, the deeper-lying
ones spiral vessels. The cellular tissue in which the
vessels are imbedded, consists of thin-walled, elongated
cells, and narrow, thick-walled cells are only met with in
the outermost parts of the ligneous body. The most pos-
terior or internal part of the vascular bundle also contains
thick- walled cells. In front of the vascular bundle lies a
bundle of liber-cells, which is connected by a process on
each side with the wood bundle ; between the wood and
the liber lies a bundle of proper vessels. It is here clear
that this vascular bundle has entirely the same structure
as the Monocotyledonous bundle, the only distinction that
occurs being, that the woody mass is gradually increased
in size by the deposition of new layers on its outside.
The relations of the vascular bundle to the surrounding
parenchyma are likewise identical with those which are
found in the Monocotyledons furnished with an outer
ring of thick-walled cells. The large thin-walled paren-
chymatous cells of the stem, which send broad processes
corresponding to the medullary rays, between the vas-
cular bundles to the bark or rind, acquire thicker walls
between the anterior portions of the vascular bundle, and
approach pretty closely in form to the liber-cells.

If we examine the young shoot of Aristolockia Sipho,
we find here also a great similarity of the vascular bun-
dles to those of the Dicotyledons,* the woody mass con-
sisting of a thin-walled parenchyma, the cells of which
do not lie in any definite order. In the back part of the
vascular bundle are found minute spiral vessels ; in front,
large porous vessels. It is only in front of these large

* Q,y. misprint for Monocotyledons. TRANS.

38 THE PALM-STEM.

vessels, after the woody mass has increased in size by
further growth, that the wood-cells begin to arrange
themselves in regular rows. In front of the wood lies
a large bundle of proper vessels, which may be readily
distinguished from the parenchyma by their thin walls,
and the absence of chlorophyll-granules. This vascular
bundle is distinguished from that of the Monocotyledons,
by the fact that the liber does not stand in immediate
connexion with the vascular bundles ; but the liber-
bundles grow together in waving lines, and are separated
from the proper vessels by several layers of cells. We
find the same structure in the vascular bundle of Meni-
spermum canadense.

Comparing the vascular bundle of the Monocotyledons
with that of the Dicotyledonous trees, we meet with a
great resemblance, the posterior portions of the wood
having the cells thin-walled, and placed in no regular
order. The smallest vessels lie farthest back, and are
spiral ; nearer to the front lie larger scalariform ducts,
and to these follow the large porous vessels. We do not
come to the firmer portion of the wood until we arrive
at the front of these, this wood being composed of thick-
walled cells lying in regular rows, and of porous vessels.

From these facts it is most clear :

1. That the vascular bundle of the Monocotyledons
exhibits the same composition, as that of the Dicotyledons
possesses in 'its earliest condition.

2. That the part of the vascular bundle of the Palms,
which I, in spite of its having no ligneous solidity, have
denominated the wood, corresponds most closely with the
innermost part of the ligneous body of the Dicotyledons,
to which Hill applied the name of the Corona.

3. That in many Dicotyledons, also, a bundle of proper
vessels lies between the liber and wood; besides the
plants named, this may be found in Spiraa Ulmaria,
Aruncus, Fumaria ojficinalis, Echinops, and Mimosa
pudica.

4. That the portion composed of prosenchymatous cells,

THE PALM-STEM. 39

lying in front of this bundle of proper vessels in the
Monocotyledons, is not to be referred to the wood, but to
be considered as the liber of these plants.

On the other hand must be mentioned, as distinctions
in these structures :

1. That in the Dicotyledons, the formation of the vas-
cular bundle is not concluded with the development of
the Corona., as in the Monocotyledons.

2. That in very few Dicotyledons is there a bundle of
proper vessels between the liber and wood.

3. That in most Dicotyledons the liber-bundle is sepa-
rated from the wood-bundle by some layers of cells; that
it does not exhibit such different degrees of development
in different parts of the bundle ; and that it never attains
to such considerable firmness and size, as in many
Monocotyledons.

It has already been observed that, besides the proper vessels, milk-vessels
are met with in many Monocotyledons ; the same occurs in many Dicotyle-
dons ; for instance, in the Umbelliferse, where milk-vessels lie among the
proper vessels in the wood-bundle, as well as in the pith and rind, while the
milk-vessels of the Dicotyledons, e. g. in Euphorbia, Asclepias, Morus, Acer,
Sambucus, &c., scarcely ever occur in the vascular bundles, but only in their
vicinity, in the bark and pith. (See Bernhardi, iiber Pflanzengefasse, p. 53 ;
Treviranus, Beitrage, p. 44; Moldenhawer, p. 126.)

Comparison of the Palm-stem with the Stem of
Dicotyledons.

So long as Desfontaines' theory, that the young vascu-
lar bundles of the Palm-stem originate in the centre, is
believed to be correct, no parallel can be drawn between
the stem of Palms and of Dicotyledons, for in such a con-
dition these stems would exhibit only distinctions, but no
resemblances. Since, however, the course of the vascular
bundle is quite different from what was formerly sup-
posed, and since its structure agrees perfectly with that
of the vascular bundle of the Dicotyledons, it appears to

40 THE PALM-STEM.

me that a comparison between the stems of the two great
classes of vegetables may be instituted without straining
the facts.

Although the vascular bundle belonging to each indi-
vidual leaf cannot be distinguished from the rest in the
cross section of a Palm-stem since the position in con-
centric circles (which should result from the similar
course taken by all the vascular bundles going to one
leaf) is variously distorted yet we may imagine the
vascular bundles going to each leaf united into a tubular
layer. This tube, however, is not cylindrical, but forms
an elongated cone. As long as the leaf which is supplied
by these bundles remains the uppermost, the conical tube,
composed of its vascular bundles, forms the most external
layer of the stem. But when a new, higher leaf is deve-
loped, the vascular bundles going to this leaf form a new
layer, which is developed on the outside of the last layer.
Since, however, the apices of these conical layers are not
closed, but the vascular bundles of which they are com-
posed run outward again from the centre of the stem to
reach the leaves, the above described crossing results,
occurring between the vascular bundles which emerge
into a leaf, and all subsequent bundles running up to
younger leaves.

If we compare with this the structure of the first year's
shoot of a Dicotyledonous tree, for excellent observations
on which we are indebted to Dupetit-Thouars (Histoire
d'un morceau de bois), we meet with a very similar
course of the vascular bundles. The vascular bundles in
it, running to the lowest leaves, which have formed the
corona on the lowest part of the branch, and which we
will designate A, pass in a curve outwards, and the
vascular bundles supplying the next succeeding leaf above
(B), lie with their lower portions on the outside of the
bundles A, up to the point where these latter turn out-
ward ; above this they form the succeeding portion of the
corona up to where they also pass outward into the leaf.
We see, therefore, that the course of the vascular bundles

THE PALM-STEM. 41

in the Palm-stem and in the one-year old shoot of the
Dicotyledons is exactly similar, and that the conception
of a different mode of growth, and the division of plants
into Endogens and Exogens formed on it, is altogether
opposed to nature.

I have always found the structure of the yearling shoot of the Dicotyledons,
such as I have described ; 1 cannot, therefore, agree with the description
given by Schweigger (Beobacht. auf naturhist. Reisen, 107), which goes to
show that in the Dicotyledons the upper leaves are supplied by the internal,
the lower by the external vascular bundles.

In spite of this resemblance, we must not overlook the
fact, that very important differences occur in the organiza-
tion and growth of Dicotyledons and Monocotyledons. In
the yearling stems of a Dicotyledon, the vascular bundles
going to the higher leaves are interpolated in their lower
part, between the liber and wood of the older. The wood
of the younger thus becomes blended with that of the
older, while in many cases the liber-bundles of these
remain isolated. From this cause, the wood in the lower
part of the stem becomes thicker, and the entire stem
acquires a conical form. In the Palms, on the contrary,
in the vascular bundles of which the liber and wood stand
in the closest connexion, the lower parts of the younger
bundles are never interposed between the wood and liber
of the old, but lie isolated in the cellular tissue of the
stem, nearer to the periphery than the older bundles.
Thus neither the liber nor wood of the older bundles
exhibit a deposition of new portions, but remain perma-
nently in that stage of development, which they have
attained at their original completion.

We must not consider it as an universal distinction, that in the Monocoty-
ledons the vascular bundles do not unite laterally into a reticulation, since,
on the one side, in Dracana, Aletris, &c., the fibrous bundles of the outer-
most layers become connected together in this way, and on the other hand,
in some of the herbaceous, and even in many woody Dicotyledons, for in-
stance, in Rosa and Rubus, this lateral connexion of the vascular bundles
does not exist.

The phenomenon that the Palm-stem grows but little

42 THE PALM-STEM.

in thickness, which at the first glance seems to indicate a
great difference of its growth from that of the Dicotyle-
dons, is readily accounted for by the smaller size of the
lower part of its vascular bundles. That it does not in-
crease in thickness at all is not a perfectly correct state-
ment, for it certainly does exhibit a slight enlargement by
growth ; this increase of diameter occurs in many Palms,
for example, in Areca oleracea and Iriartea ventricosa,
not so much at the inferior extremity as higher up in the
stem, whence this acquires a spindle-like form. Since
the lower part of the bundles is not thicker than a hair,
it will be conceivable how many thousands of them may
be formed beneath the rind of the stem, without increas-
ing its diameter more than a single inch, which is so
slight a growth that it is wholly overlooked.

I have already stated that the fibrous bundles in Dra-
caena become more strongly developed, and that the stem
then grows in thickness as in the Dicotyledons. From
the circumstance that these bundles, lying in the ex-
ternal layers of Dracaena, are nothing else but lower
extremities of the vascular bundles of the stem, it is most
evident that their growth and development is not to be
regarded as anything different from the growth of the
apex and the centre ; and that the idea of a double growth,
which Mirbel (Annal. du Museum, xiii, 67) imagined
that he found in Dracaena, Yucca, Aloe, Ruscus, Smilax,
Dioscorea, and Tamus, is no less incorrect than the idea
of the central vegetation generally attributed to the
Monocotyledons.

Obs. 1. I have mentioned above that Moldenhawer had already expressed
himself against the correctness of the view of Desfontaines. Moldenhawer
distinguished in the same way, as has been done above, in every vascular
bundle of Zea Mays and Bambusa, in which plants his investigations were
chiefly made, liber, proper vessels, and wood; he also found that, in the
Grasses and Palms, the younger leaves were supplied by the outer, the older
by the inner vascular bundles, (Beitrage, p. 50.)

Thus far, therefore, our researches fully accord ; but they differ in many
points in respect to the fibrous bundles devoid of vessels, and the origin of
the wood, the reason of which doubtless lies in the fact, that Moldenhawer

THE PALM-STEM. 43

neglected to examine the same vascular bundle in various parts of its course,
so that its changes of form and structure remained unknown to him.
Moldenhawer states that both in the Grasses and the Palms, fibrous bundles
without vessels are formed beneath the rind, and believed that shortly after
their formation proper vessels, and subsequently the wood portion, were de-
posited on their inner side, and that in this way the fibrous bundles became
vascular bundles. I cannot confirm this account, for the transition from the
evascular fibrous bundle into the vascular bundle is merely one relating to
space, and has no reference to its development in time ; the lower part of
every vascular bundle remains permanently in the condition of an evascular
bundle of prosenchymatous cells, while its upper portion does not appear,
even in its earliest stage, in the form of a liber-bundle ; but even at a period
when it is still of a gelatinous delicacy of consistence, the rudiments of all
the parts which it subsequently contains may be detected in it.

Moldenhawer further states, that in the Palms, not only has each vascular
bundle its proper liber, but there exists besides a general liber, lying under
the rind (1. c. p. 56). There is found, he says, in Phwnix and other Palms,
a line of separation between the wood-layer and the rind, from which those
fibrous bundles develope inwards, which are changed into vascular bundles,
but on its outside, those which contain only fibrous cells, the latter being
comparable to the liber of trees. A correctly observed fact certainly appears
to afford the ground for this opinion. I have observed already, that in many
Palms, especially in the Cocos-like stems, the rind becomes gradually thickened,
and this in an irregular manner ; that larger or smaller portions of the cel-
lular tissue in which the fibrous bundles are imbedded acquire thickness in
their walls, and form a firm, apparently dead envelope. In this change of
the outer parts of the fibrous layer, the newly-developing fibres cannot be
produced on the outside of the older ; they must, therefore, originate in the
interior of the fibrous layer, and the matter may thus acquire some resem-
blance to the formation of liber in the Dicotyledons. But a second circum-
stance also has to be considered, which might have contributed still more
readily to render Moldenhawer's opinion, that the Palms possess a general
liber, plausible. In those species, namely, where, as in Cocos, the outer
fibrous layer is very thick, in tracing their evascular fibrous bundles, it is
found that they do not all enter into the interior of the stem, and become
converted into vascular bundles, but that a portion of them pass immediately
into the petiole. But as those fibrous bundles are changed, like the rest,
into vascular bundles in the petiole, they cannot be regarded as liber-bundles
corresponding to the liber of the Dicotyledons.

2. Another circumstance remains to be mentioned, occurring in many
Palms, and which at first sight does not appear to harmonize with the theories
of their growth detailed above. We find in many Palms, that their evas-
cular, small, fibrous bundles do not all lie between the rind and the developed

44 THE PALM-STEM.

vascular bundles, but some of them are met with scattered among the vas-
cular bundles of the stem, for example, in Astrocaryum vulgare, Cocos botry-
ophora, coronata, Leopoldinia pulchra. I have met with this modification
principally in the Cocos-\\ke, stems, in others only in rare cases. These
fibres, therefore, only occur in such stems as possess a fibrous layer abun-
dantly furnished with fibres, and a great many vascular bundles. Consequently
it is probable, that in the great mass of closely-crowded bundles which fill
up these stems, circumstances readily arise, which prevent the nascent fibres
from becoming developed in the normal positions, and give rise to their
origin in unusual places. These scattered fibres, too, do not seem to occur
in all specimens of the same species, at least they were very abundant in one
stem of Leopoldinia pulchra, while in a second they were altogether wanting,
which evidently proves that accidental causes give occasion to their origin.

3. Mirbel fell into still greater error than Moldenhawer in his researches
into the course of development of the vascular bundles of the Monocotyledons.
He states (Annal. du Museum, xiii, 69) that each vascular bundle is formed,
at its origin, solely of one bundle of large ducts (reticulated vessels), that
around these a tissue of delicate tubes is gradually formed, by which is un-
derstood the cellular tissue of the wood, the spiral vessels, annular vessels,
the proper vessels, and the liber-bundle, these parts not being distinguished.
The membrane of these tubes, then, gradually becomes so much thickened
that at length their cavities are filled up.

4. In a description of the vascular bundle of Calamus (Ann. d. Sc. nat. ii,
pp. 229-236), Amici attributes a different import to the different parts of
the vascular bundle from that which I have done. Amici considers the
wood-cells to be liber-tubes, the liber-tubes proper vessels, the proper
vessels he indeed recognises as thin-walled and not porous tubes, but leaves
it undetermined to what kind of organ they belong. The description and
representation of the vascular bundles of Calamus, given by Kieser (Phytot.
p. 121, Tab. iii, fig. 29), deviates still more from the truth, since the liber-
and wood-layers are not distinguished at all, and the proper vessels are
regarded as spiral vessels.

5. I shall readily be excused from detailing minutely the views of
Lestiboudois ; he believes that the Palm-stem is to be compared with the
bark of the Dicotyledons, he finds nothing analogous to wood and pith.
(Principes de Botanique, pp. 149-158.) This strikingly testifies how little
he is acquainted with the minute anatomy of plants.

THE PALM-ROOT. 45

OF THE ROOT OF THE PALMS.
Form of the Root.

The full-grown Palm-stem is never furnished with a
tap-root, but its inferior portion, rounded off like a bulb,
is clothed with a quantity of fibrous, branching roots.
These roots are always slender, not very long, beset in
an irregular manner with slender lateral branches, cylin-
drical, obtuse at the extremities, in the young condition
white, and subsequently brownish. Their young shoots
are clothed with fine hairs ; sometimes short spinous
elevations occur on the root, which are to be regarded as
abortive lateral branches. The germinating Palm has a
tap-root, but this attains no considerable size. Shortly
after germination, side roots are developed from the base
of the stem ; the tap-root disappears, and after a time
the first side roots also die away, and are replaced by
new rootlets, which spring out in a circle above the earlier
ones. This process is repeated in a manner analogous
to that in the bulbous plants, Although the roots arise
very closely crowded together, yet the subterraneous por-
tion of the stem soon becomes completely covered with
roots, and the new ones then arise, as in Pandanus,
above ground. In this way it often happens that the
stem stands raised free above the surface of the earth,
merely supported by the roots ; for example, in Iriartea
exorhiza.

The formation and first development of the roots occur
in the interior of the stem, between the fibrous layer and
the developed vascular bundles. In this place is formed
a nucleus of cellular tissue (a true bud), which acquires
the shape of a root, and breaks through the rind. Such
buds of future roots may be found in considerable abun-

46 THE PALM-ROOT.

dance for the distance of some inches above the youngest
circle of roots, when the bark and fibrous layer are cut
open down to the vascular bundles.

Anatomical Examination of the Hoot.

The Palm-root is composed of two distinctly separated
layers an outer, looser, and spongy cortical substance,
and a tough, woody, central bundle.

The cortical substance is coated by a parchment-like
membrane, under which lies a white spongy mass, in
which in some species lie liberous fibres ; in others these
are wholly wanting. Toward the extremity of the root,
and in the young side roots, this cortical layer is suc-
culent and compact ; in the older parts of the root it is
often half dried and more lax. The central bundle is
formed of a compact woody substance, which cannot be
separated into distinct individual bundles like the wood
of the stem. The central bundle of the side roots is
immediately connected with that of the main roots.

When a root is traced backwards into the stem, it is
found that the cortical part of the root is considerably
diminished at its entrance into the cortical layer of the
stem, after a short distance disappearing by blending with
the cellular tissue of the stem. The central bundle, on
the contrary, penetrates the fibrous layer of the stem, and
spreads out on the outer layers of the wood-bundles of
the latter in the form of a disk. In its way through the
fibrous layer, it already begins to divide into several bun-
dles, separated by thin layers of cellular tissue. When
it arrives at the woody layer of the stem, it divides into
a great number of delicate filamentous bundles, which
run out in all directions like the rays of a star, and,
passing in serpentine course between the vascular bundles
of the stem, enter into its interior. They cannot be
traced more than about half an inch down between the
woody bundles, because they continually divide into

THE PALM-ROOT. 47

successively finer portions, apply themselves upon the vas-
cular bundles of the stem, and there terminate.

Besides this connexion with the interior of the stem,
the root is most intimately connected with fibrous layer
by means of its cortical portion; for, into the cortical
portion of each root penetrates a portion of the evascular
fibrous bundles of the stem. In some roots, as in those
of Cocos and Plicenix, the little fibrous bundles run still
further out into the roots, and then lose themselves gra-
dually ; in others, as in Diplothemium maritimum, Sabal
Adansonii, they are at once lost at the very beginning of
the root.

In Dracaena Draco., also, I observed that the outer, fibrous, firm layer of the
stem penetrated the roots, was continued some distance in this, and formed
a sheath round its central body, which was far stronger and longer on the
side turned towards the surface of the earth than on the under. This
fibrous layer was gradually attenuated, and altogether lost in the distance
of a few inches. It is clear from this, that the opinion of Dupetit-Thouars,
that the roots are formed of the fibres running down from the leaves and
buds, is totally erroneous.

The roots of different Palms possess a very similar
organization. That of Diplothemium maritimum may
serve as an example. In the cross section, it is seen that
in the central bundle all the vessels lie toward the cir-
cumference, and the middle is formed solely of cells.
The vessels are constantly placed in such a manner that
the largest are nearest the centre, the smallest nearest the
circumference ; therefore the condition is the contrary of
that which we have met with in the stem. The vessels
do not lie, as in the vascular bundles of the stem, irre-
gularly scattered and isolated, but in rows, which are
directed from the centre toward the circumference, and
these rows are frequently split toward the exterior into
two diverging arms. The largest of these vessels present
the form of reticulated vessels, and are composed of rather
short tubes, possessing reticularly perforated septa at their
ends. The small vessels, situated more externally, ex-
hibit the form of porous and scalariform ducts. In

48 THE PALM-ROOT.

respect to the epoch of their development, the latter agree
with the spiral vessels of the vascular bundle of the stem,
for they are already completely formed at a time when
the large vessels still appear as thin -walled tubes without
fibrous deposits, and the cells of the root have yet but
extremely delicate membranes. The vessels are sur-
rounded by rather thick-walled, much elongated cells,
with horizontal septa. But only those cells situated next
to the vessels exhibit these horizontal septa, and they pass,
in the interspaces between the vessels and in the central
space inclosed by the vessels, into prosenchymatous cells,
which, in the centre of the vascular bundle, are again
transformed into elongated parenchyma, and form an
analogue to the pith. The whole central body is sur-
rounded at its circumference by a few layers of thin- walled
prosenchymatous cells, to which follows a row of narrow,
elongated thick- walled cells.

Between every two groups of vessels lies a bundle of
proper vessels ; the size of this is ruled by the length of
the rows of vessels lying beside it, for only a small roundish
bundle lies between the short rows, while those situated
between the long rows extend far in toward the centre of
the root. The arrangement of the proper vessels is always
such, that those lying internally are very wide, the outer
very narrow : the narrower and wider are not, as in the
stem, mingled together. The membranes of these vessels
are always thin, not unfrequently finely porous, which
also sometimes occurs in those of the stem of Tamus
elepliantipes. They are composed, both the wide and
narrow, of closed elongated tubes, with horizontal or
diagonal septa; like those of the stem, they contain an
opaque, granular sap.

The cortical substance consists chiefly of regular thin-
walled parenchyma-cells, with intercellular passages.
Those cells in the vicinity of the vascular bundle have no
intercellular passages between them, and are somewhat
expanded in the direction of their breadth. The inter-
cellular passages also disappear toward the surface of the

THE PALM-ROOT. 49

rind ; the cells become larger ; the outer, at the same
time, acquire thickness in their walls, and form the coria-
ceous membrane above described. The epidermis is
composed of cells not elongated, but papilliform and
projecting. In the young root the rind is compact ; but
with the greater expansion of the root in thickness, and
the decrease of its juices, the cells separate from each
other in isolated, larger or smaller, irregularly scattered
places, and thus form irregularly distributed air-cavities.
The cells immediately surrounding the central body, con-
tain on their inner side transverse fibrous thickenings,
like many anther-cells. Isolated cells, scattered without
order in the rind, contain raphides.

The lateral roots spring from the larger roots in the
same way as these do from the stem. A nucleus of cellular
tissue is formed between the rind and central body of
the main root, and in this are developed moniliform
vessels. On further enlargement, this nucleus breaks
through the rind of the root, and appears as a branch of
it. The moniliform vessels penetrate the central body of
the root, run in the most varied directions in the cellular
tissue situated between the vessels, and finally apply
themselves to the sides of the large vessels. In the lateral
root itself, the vascular tubes become more and more
elongated, and form, with a portion of the cellular tissue
which they inclose, and with the elongated cells by which
they are surrounded, the central body of the root-branch.
The most external layer of these elongated cells is blended
with the thick-walled cells surrounding the central body
of the main root ; while the more central, pith-like cells
of the lateral root are connected with the parenchyma
situated beneath these elongated cells. The rind of the
side root is, indeed, separated from these in the greater
part of its course through the rind of the main root, by
its epidermis and its outer layer of elongated cells ; but
both the elongated and the parenchymatous cells of the
rind of the side root are blended with the more internal
layers of the rind of the main root.

50 THE PALM-ROOT:

The connexion of the root with the stem takes place in
a manner wholly analogous; since in this also, in the
dissolution of the central body into single fibres, its
vessels are distributed into a great number of fine moni-
liform vessels. Those fibres penetrating the stem are,
indeed, formed for the most part of thin moniliform
vessels; but they nevertheless present an organization
tolerably resembling that of the vascular bundle of the
stem, for their vessels are surrounded by cellular tissue,
a little liber-bundle lies upon their outer side, and their
proper vessels occur between this liber-bundle and the
wood.

I found the structure described, perfectly similar in the roots of a con-
siderable number of Palms, as might indeed be expected, since the said struc-
ture occurs almost as universally in the roots of Monocotyledons as the
above-described structure of the vascular bundles in their stems.

This structure, however, is not without exceptions. I have already said
that in the upper parts of many Palm-roots, e. g. Phoenix, Cocos, fibrous
bundles are scattered through the rind, while no trace of them is found in
others. But the rather thick root of Iriartea exorUza exhibits more im-
portant deviations. A cross section of it presents to the naked eye a star
composed of brown lines, with obtuse, mostly bifid, rays. The microscope
shows that this star is formed of crowded vascular bundles. Besides these,
scattered vascular bundles occur singly in the centre of the star, but a central
cord, like that in the other Palm-roots, is wanting. The vascular bundles
have a structure totally unlike that of the vascular bundles of the stem, the
relative position of their organic systems being quite different. The liber-
bundle is directed towards the centre of the root, and in some of the vascular
bundles lying in the middle of the root, surrounds the woody portion. The
liber-tubes have thicker walls, and are more numerous in the outer bundles
than in those situated in the middle. The woody portion has 1, in rare
cases 2-3 large vessels, which in the outer bundles are &-& in the
inner, 77-73 f a line i n diameter. They are composed of longish tubes,
some have horizontal septa, some are without these ; their walls are covered
with little pores, placed in longitudinal rows ; where two vessels are in con-
tact, they have the form of scalariform ducts. They are surrounded merely
by 1-2 rows of parenchymatous cells. The outer side of this woody portion
is in contact with the parenchyma of the root. The bundles of proper vessels,
of which each vascular bundle contains one or two groups, consist, as in the
stem, of mingled, wide and narrow tubes. These do not lie between the
wood and liber, but in most cases on the outside of the liber-layer, half

APPENDIX. 51

imbedded in it ; in the inner vascular bundles, the proper vessels are some-
times wholly surrounded by the liber-tubes. The outer vascular bundles
are often confluent in pairs, in which case the compound bundle possesses
four groups of proper vessels, or, from the confluence of one pair, frequently
only three.

The parenchyma of the root consists of thin-walled, somewhat elongated,
porous, parenchymatous cells, which in many places leave irregular spaces
between them. Around the stars formed by the vascular bundles, separated
from them by a few rows of cells, runs a darkish line, formed by cells with
rather thicker walls. Scattered through the whole root, and accumulated in
especial quantity in its outer layers, lie evascular, fibrous bundles, the outer
of which are often blended together, and are composed of thick-walled, prosen-
chymatous cells ; in the middle of those situated more internally, occur from
one to two thin-walled cells, which probably belong to the system of proper
vessels. I can state nothing concerning the connexion of these fibrous
bundles with the stem, as I had no opportunity of examining the latter.
These roots of Iriartea are clothed with short thorns, which are to be looked
upon as abortive lateral branches, and have their origin from the vascular
bundles of the main roots.

Different as this root appears to be from those of other Palms, yet it is clear
that its organization agrees with that of the latter in many respects, since
most of the characters by which the vascular bundles of the roots are dis-
tinguished from those of the stem, occur in the root of Iriartea, as well as
in the roots of other Palms. The reversed position, which is exhibited by
the vascular bundles of the roots of Iriartea, corresponds to the circum-
stance met with in other Palm-roots, that the cells lying behind their
vessels likewise have the form of prosenchymatous cells. In both varieties
of the root the large vessels lie furthest inward, the small ones externally ; in
both, the proper vessels form special bundles not inclosed between the wood
and liber.

APPENDIX.

It will not be unfitting to append to the preceding
description of the Palm-stem, a short account of the
works on this subject, which have appeared since it was
first published.*

* My purpose here is not an enumeration of all the observations on this
subject which have appeared since my Anatomy of Palms, but only an

52 THE PALM-STEM:

Pre-eminently deserving of mention is the acute Essay
of Meneghini (Ricerche sulla Struttura del Caule nelle
Piante Monocotiledoni, Padova, 1836), in which the
structure of the stem of Monocotyledons is traced in its
various forms, and described with great clearness. So
far as this exposition relates to the arborescent Monoco-
tyledons, the following points might indeed be brought
forward in opposition to my description. Meneghini,
who, in his description of the course of the vascular bundle,
traces it as I did from above downwards, devotes especial
attention to the changes which its position undergoes
during the development of the terminal bud into stem,
and, in reference to this, lays particular stress upon the
circumstance that, so long as the leaves remain inclosed
in the bud, only that part of the vascular bundle exists,
which in the full-grown bundle runs from the middle
of the stem downwards and outwards, and that its upper
portion is first developed with the development of the bud
into stem, and with the emergence of the leaves from the
centre of the bud out to the lateral surface of the stem.
From this passage outward of the leaf from the centre of
the bud, is derived the curve of the vascular bundle in the
centre of the stem, and the outward direction of the course
of the upper part of the vascular bundle. Meneghini paid
especial attention to the examination of the circumstance
that the vascular bundles present, not only this curve out-
ward, but at the same time a lateral bend, so that their
lower extremity does not lie in a perpendicular direction be-
neath the upper end, but diverges to its left or right. I also
had observed this circumstance in the Palm- stems I inves-
tigated, but attached no importance to it, as I considered
the oblique course of the fibres an accidental deviation.

examination of such works as have had an influence on the progress of the
study of the structure of the stem of Monocotyledons, and especially of the
Palms, whether by the announcement of new, hitherto unnoticed facts, or by
the extension of the study in its theoretical aspect ; I therefore shall pass
over, in particular, Gaudichaud's works, since the researches on which his
theory rests, have too much the stamp of superficiality to allow me to expect
any profit from them to the study of the structure of plants.

APPENDIX. 53

Meneghini has shown, on the contrary, that this condition
occurs in all Monocotyledons ; for the elucidation of this
he did not select Palms, but principally the stems of
Dractena, Aletris, and Yucca, in which, however, if his
theory be correct, the oblique course of the fibres de-
viates materially from that occurring in the Palms. In
order to give a review of the opinion promulgated by
Meneghini, an examination of his investigation of Dra-
caena and the allied forms is necessary. Meneghini states,
in regard to Dracana Draco, that no fibre of its stem runs
vertically, but that the upper part proceeds inward in the
direction of a radius, toward the centre of the stem ; near
this it turns downwards, and at the same time to the right
or left, and then runs obliquely downwards and outwards,
so that its inferior extremity lies beneath the surface of
the stem,, to the right or to the left of the upper end.
Perhaps no two vascular bundles of the same leaf exhibit
the same course, for some penetrate almost to the centre
of the stem before they turn downwards, others acquire a
curve at a short distance from the rind, some deviating to
the right, others to the left of the perpendicular line.

The changes which the leaf undergoes during its deve-
lopment are regarded as the cause of this oblique course
of the fibres. As the adoption of this point of view by
Meneghini is wholly peculiar, and forms the most essential
part of his doctrine of the structure of the Monocotyledo-
nous stem, we must enter somewhat more minutely into it.
Meneghini assumes that, in all the Monocotyledons, the leaf
originates in the form of a closed, reversed funnel, which
subsequently becomes torn up along one side, either the
whole way or only at its upper part, by the pressure of the
younger leaves succeeding it in the interior. When only
the upper portion is torn, the lower part forms the sheath
of the leaf, and the leaf remains perfectly amplexicaul ; in
the other case, the relation of the base of the leaf to the
circumference of the stem does not necessarily remain
the same in its further development, but the stem usually
grows, proportionately, more in thickness, than the base

54 THE PALM-STEM:

of the leaf in breadth, and thus the leaf occupies so much
the smaller arc of the circumference of the stem, the older
the latter grows ; while in other cases, the converse may
take place, the leaf grow broader than the stem, and the
two borders of the leaf overlap. As examples of such
plants, in which the originally perfectly amplexicaul leaf
only partly embraces the stem when full-grown, Aletris
fragrans and Dracaena Draco are, in particular, brought
forward ; in the former, the full-grown leaf embraces f of
the stem, in the latter, the leaves at the apex of the year-
ling-shoot are still perfectly amplexicaul, while on older
stems the cicatrices of the leaves only extend i round
the stem. This proportionately greater expansion of the
stem in thickness is connected, in Dracaena, with an
abbreviation of the already perfectly formed internodes,
as is apparent in a comparison of the length of the inter-
nodes of the yearling- shoot with those of the old stem,
and as even a superficial examination of the growth of
Dracaena shows ; for the rapid growth of the yearling-
shoot does not by any means correspond to a propor-
tionate elongation of the whole plant (p. 22). A necessary
consequence of this change of the relative proportion of
the breadth of the leaf to the thickness of the stem is, a
change of the original position of the vascular bundles in
the interior of the stem. So long as the young leaf sur-
rounds the stem and lies in the middle of the bud, the
vascular bundles (corresponding to the lower portions of
the complete vascular bundles) run from the whole peri-
phery of the stem, in a radial direction, towards the base
of the leaf. As the bud unfolds, the internode per-
fected beneath the leaf to which it belongs, and the leaf
proceeds outwards from the centre of the bud towards
the periphery of the stem, the upper portions of the vas-
cular bundles are developed, which, in consequence of this
outward movement of the leaf, assume a curve from the
centre of the stem towards the exterior, and run out in
the direction of a radius. Since, however, during the
completion of the leaf, the proportion of its breadth to

APPENDIX. 55

the circumference of the stem diminishes, the upper part of
the vascular bundles must also follow the leaf, and, there-
fore, those vascular bundles which enter the sides of the leaf,
instead of being directed in a radial direction to all sides
of the stem, as in the perfectly amplexicaul leaf, are only
curved towards the arc of the circumference of the stem,
on which the leaf is inserted, and since the lower part of
each vascular bundle retains its position more or less un-
changed, the result is the described condition of a double
curvature, downwards and to the side. The greater the
difference between the circumference of the stem and the
breadth of the base of the leaf, the shorter the internodes,
the greater must be this divergence ; hence it is greater in
Dracaena than in Yucca,iu Yucca it is greater than iiiAletris.
Before I go further into the detail of Meneghini's
theory, it will be advantageous to test the facts on which
it rests. In this I pass over the question whether the
leaves do originate under the form of closed funnels, or
not, as this has no essential influence on the theory. The
only essential question is, whether or not the breadth of
the base of the leaf does gradually diminish its proportion
to the circumference of the stem with the age of the latter.
That such a change must lead to the alteration in the
course of the vascular bundle which Meneghini has
deduced from it, does not admit of a doubt ; at all events,
if it be presupposed that at the time when this change
takes place, the vascular bundles running to the younger
leaves, which cross those of the leaf that has begun to
change its position, do not oppose any hinderance to the
movement of the latter. I believe that this certainly
must be the case, and this circumstance alone already
makes the theory of the Paduan phytotomist appear
untenable ; if, however, we even overlook this, there exist
other, more definite reasons, which show that the unequal
growth of the leaf and the circumference of the stem
do not occur in the way Meneghini has represented.
According to his statements, this process is most clearly
to be seen in Dracaena Draco, for the leaves, or their

56 THE PALM-STEM:

cicatrices, embrace a smaller portion of the circumference
of the stem, in proportion as they are situated lower
down upon it, so that the cicatrices only occupy a third
of the lower part of the stem, while the leaves at the
summit of the yearling- shoot completely surround it.

I can only confirm the smallest possible part of this
statement. I decidedly found no change of the relative
position and size of the cicatrices, in the stems which I
examined in reference to Meneghini's statements, (the
largest being about twenty-seven feet high, and thirty-
seven centimeters in circumference at the lower part), but
the scars of the leaves just fallen exhibited the same pro-
portion to the circumference of the stem as the scars at
the lower part of the stem, i. e. they extended round
about -| to J- of the stem. Measurements, which I insti-
tuted on different cicatrices, showed that small variations
occur in the proportion of the breadth of the scar to the
circumference of the stem, which are not at all connected
with the age of the stem. Removal of the leaves from
the terminal bud showed, that not only is the same pro-
portion of the base of the leaf to the circumference of the
stern to be found in the leaves which are seated on the
conical portion of the stem, concealed in the leaf-bud, but
that it also holds for those leaves situated on the upper,
flattened surface of the axis of the bud, excepting the
innermost. As, on the other hand, continuing the re-
moval of the leaves, we approach the youngest leaflets,
this proportion alters, for the borders now certainly do
approach each other. In one bud I examined, the borders
of the innermost leaflet in which the lamina was distinctly
formed, were approached to within i of the circumference
of the circle ; the next leaflet appeared in the form of a
perfectly amplexicaul cone, about one millimeter in dia-
meter, at the base of which was to be seen a narrow,
short slit ; it had exactly the form of the cotyledon of a
Monocotyledonous embryo ; in the succeeding leaflet, the
slit formed by the borders of the leaf was again more
widely opened. This observation does indeed fully con-

APPENDIX. 57

firm the assumption of Meneghini, that in Draccena Draco
the perfectly amplexicaul leaf is changed into one which
does not surround the stem, but an essential distinction
exists, in that Meneghini states this change to begin after
the completion of the leaf, and to be continued further in
the cicatrix after the fall of the leaf; while the result of
my investigation is, that the leaf only surrounds the stem
so long as the part of the axis of the bud on which it is
inserted is still in a rudimentary condition, and the thick-
ness has not, or scarcely, exceeded a millimeter; that in the
next older leaflet the borders are already \ of the cir-
cumference of the stem apart, and the permanent propor-
tion between the base of the leaf and the circumference
of the stem already exists in the leaflets situated but a
couple of internodes further out. This difference in the
results of our investigations is very much the more im-
portant, as will be shown further on, in so far that such an
alteration of the proportion of the breadth of the base of
the leaf and the circumference of the developed stem
must cause extraordinarily great changes, which we never
see, in the internal structure of the stem, and in its
growth in thickness ; while such changes, if they begin in
a portion of the stem a millimeter thick, and cease in a
part only a few millimeters in diameter, take place in a
part which consists merely of soft cellular tissue in full
process of multiplication, which exhibits extremely rapid
growth in all its parts, in which, therefore, the softness
and continual metamorphosis of its substance, present no
mechanical hinderance to a more rapid development of
one part, and a less extensive development of another.
That these changes of form exert influence on the lateral
divergence of the vascular bundles cannot, indeed, be
denied ; but, on the other hand I cannot think, that they
are the sole, or even the principal cause of their oblique
course, inasmuch as this occurs in the vascular bundles
of Aletris fragrans, although no trace of an alteration of
proportion between the base of the leaf and the circum-
ference of the stem is to be discovered here.

58 THE PALM- STEM:

According to Meneghini's statements, the borders of
the leaves should here also separate so far, that the cica-
trix only surrounds of the stem; but I cannot give
the least confirmation to this statement. The borders
of the leaf overlap a little at the base ; this condition is
no longer visible on the cicatrix after the leaf has fallen,
for this appears simply encircling the stem. However,
this peculiarity is exhibited not merely by the young
stems, from which the leaves have newly fallen, but also
by old stems ; and, in particular, on a stem thirty-eight
centimeters in circumference, in the Tubingen Garden,
the leaf-scars are perfectly evident, and still exactly sur-
round the stem, although, from the great increase in
thickness, its rind presents many longitudinal fissures.
Consequently, observation of the surface of the stem, de-
cidedly does not confirm Meneghini's view as to the
alteration of the leaf of Aletris, and the subsequent
changes in Draccena ; and the matter might here have
been regarded as settled, but that the objection that in
other specimens than those which I have examined, the
conditions might exist such as my honoured friend has
described, rendered it to the purpose, to institute a more
minute examination of the changes which a stem would
necessarily undergo, the circumference of which became
more enlarged than the bases of the leaves, in the way
described, after its leaves were perfectly formed or had
fallen, since this examination will show that this process
cannot possibly present itself in stems with abbreviated
internodes. I have before me a portion of a stem of
Aletris fragrans 31 inches long ; at the upper end it is
10' 5 lines in diameter, at the lower 13 lines; the ex-
ternal, hard, fibrous layer is 1 line thick above, 2' 5 lines
thick below: there are upon it 57 leaf- scars, hence an
internode is, on an average, 6*5 lines long. The cica-
trices surround the stem, and are placed obliquely, in such
a way, that one part of each of them, corresponding to
the middle line of the leaf, is lower down than the oppo-
site part corresponding to the borders of the leaf; so that

APPENDIX. 59

in each case the point of insertion of the borders of a
leaf is approximated to within about three lines of the
point of insertion of the central portion of the leaf next
above it. If, now, we imagine the lowest internode to
expand so much in breadth (without enlargement of the
cicatrix), that the meeting borders of the cicatrix would
be separated about \ of the breadth of the cicatrix, a
portion of stem, nearly eight lines in breadth, must be
inserted between the separating borders of the leaf; and
during the period of the completion of the structure of
this piece of stem, new tissue must be formed in the in-
terior; the firm, fibrous layer, already 2*5 lines thick,
must enlarge a fifth part of its circumference ; and the
inner soft substances must increase in quantity, in like
proportion, to fill up the space within this expanded
layer. Of all this, no trace can be found in the examina-
tion of a stem of Aletris. Instead of what this theory
requires, the insertion of an entire segment of a circle
between the old parts, and the enlargement of the in-
ternal soft substance, we find that the latter, when once
formed, remains for ever unchanged, and in like manner
no new development is met with, in any part of the sub-
stance, of firm woody layers ; but we do find that new
fibres and cells are produced uniformly in a true cam-
bium layer around it, and upon its outside. This sta-
tionary condition of the internal parts of the stem, is
again, fully sufficient of itself to refute the whole of
Meneghini's theory. But if, disregarding this fact, we
imagine the lowest internode, the cicatrix of which in the
stem under consideration was 39 lines in circumference,
to undergo the change described, this would obtain,
by the addition of \ of the size of the base of the leaf,
a circumference of 47 lines. In this expansion the
next internode above must share^ since we always find
the stem of Aletris to be nearly cylindrical. But the
part of the leaf next above, which lies only a couple of
lines to one side, is situated only about three lines above
the borders of the lowest leaf, from which we set out.

60 THE PALM-STEM:

With such a short distance between the lower and upper
leaf, the expansion of the lower intern ode must be con-
tinued into the part of the upper leaf which stands ver-
tically above the growing portion of the lower internode ;
since the fibres, which would form the piece of stem inserted
between the borders of the leaf of the lower internode,
must, like all other fibres of the plant, be continued in a
tolerably straight direction upwards in the stem ; hence
the cicatrix of the second leaf must become wider. The
second cicatrix, in the stem in question, is likewise 39 lines
broad, but we must add about eight lines for the expansion
of the lower internode ; therefore, we obtain 47 lines as
its breadth. If we next suppose, in accordance with
Meneghini, the growth in like manner of a piece i of
the breadth of the leaf between the borders of the leaf of
of the second internode, the circumference of this inter-
node would be increased at least nine lines, and we thus
obtain a circuit of 56 lines. This internode would, con-
sequently, exceed the lower about nine lines in circum-
ference. Since we do not find this in nature, and since
we must assume that the fibres which grow between the
borders of the leaf of the second internode must be con-
tinuous into the lower internode, we are compelled to
assume that the lower internode shares in the expansion
of the upper, and, consequently, also acquires a circum-
ference of 56 lines. But if, now, that which occurred
in the second leaf hold also exactly for the third, its
middle portion must expand about nine lines in breadth,
because a piece of stem equal to one sixth of 56 has
grown beneath it, in the internode of the second leaf.
The breadth of the second leaf amounted to 47 lines ; the
third would, therefore, be 56 lines broad. Then allowing
the borders of the leaf to separate about J of the breadth
of it, we obtain 67 lines for the circumference of the in-
ternode. In this enlargement, the lower internodes again
naturally take part; we have, therefore, raised the circum-
ference of the stem from 39 to 67 lines by these changes
of only three internodes, and yet we are still far below

APPENDIX. 61

the size which the stem must acquire, since this expan-
sion of the lower intern odes would again be followed by
an expansion of their cicatrices in breadth, which would
again necessitate an increased expansion of the part of
the stem lying between the borders of the leaves (for this
part should equal J of the cicatrix) ; and in consequence
of this, the upper leaves would be again expanded in
breadth, which expansion would react again upon the
lower internode, and so on. It is easy to see that this
cannot be the proceeding which nature follows in the
growth of these stems, since it would lead to an immense
increase of thickness, even if it were restricted to a con-
dition where the margins of the leaves were separated
about i of the breadth of the leaf at the complete develop-
ment of the latter; and that in Draccena, where the
separation of the margins of the leaves would equal
double the breadth of the latter, the increase of the dia-
meter of the stem must occur in a far more excessive
proportion. Since, therefore, on the one hand, these
conclusions, which are necessary deductions from the pro-
cesses asserted by Meneghini to occur, would lead to
impossibility, and, on the other hand, both direct obser-
vation of the cicatrices and the anatomical examination
of the stem testify most clearly against this notion of an
unequal growth of the leaf-scars and the circumference of
the stem, the divergence of the fibres from the perpen-
dicular line cannot be caused by the mechanical process
supposed by Meneghini.

Even if this oblique direction of the fibres were a con-
sequence of the unequal growth of the leaf-scar and the
circumference of the stem in Dracana, Aletris, &c., this
derivation of it would not be applicable to the Palm-stem,
since this always has its leaves and leaf-scars surrounding
the stem. Meneghini did not overlook this circumstance ;
but he imagined he had discovered a second mechanical
cause, which produced an oblique position of the fibres
also in the stems with completely amplexicaul leaves, and
which consisted in a movement of the leaves to one side.

62 THE PALM-STEM:

He starts from the fact, that the leaves of the arborescent
Monocotyledons, e. g. of Yucca, lie in a heliacal line,
which in the flattened bud of these plants passes into a
true spiral. The heliacal line marks a constant relation ;
this does not exist in the spiral, for this passes, with the
further development of the leaves lying in it, and the
conversion of the bud into stem, into the heliacal line
running upon the side of the stem ; in this, the same
lines still remain, since they always gain at their inner
end as much as they lose at their outer end by passing
into the form of a helix. Each leaf originates in the
centre of the bud, and passes, when a new leaf arises
in the centre, into the place of the preceding one, till
at last it comes to be situated on the outer surface of
the stem. In this movement each leaf follows a spiral
course, in which the mutual relations of the leaves remain
the same, and all the leaves are carried back uniformly.
Each leaf therefore exhibits, besides the movement from
within towards the outside, and from above downwards,
also a horizontal motion in a spiral direction ; on the first
of these motions depends the divergence of the vascular
bundle from the vertical line ; the second gives rise to a
divergence in the horizontal direction, since the upper
part of the vascular bundle, running from the centre of
the stem to the leaf, follows the lateral motion of the
latter. Since the turns of the spiral are closer in the
middle, and the motion of the leaves becomes slower in
proportion as they pass further out, the greatest bending
of the vascular bundles takes place in the centre of the
stem (1. c. p. 16.)

Against the idea that, in the development of the ter-
minal bud into stem, the leaf-spiral passes into a helix,
and that each leaf traverses the length of this spiral, no
objection can be urged ; this traverse of the spiral, how-
ever, is only a seeming movement, and by no means con-
nected with an actual motion to the side. If Meneghini's
idea were correct, it is clear that the most external leaf
seated on the spiral, could not advance with it in a lateral

APPENDIX. 63

movement towards the surface of the stem, without the
divergence between it and the uppermost leaf upon the
helix becoming diminished, until the leaf had arrived at
the lateral surface of the stem, and there become perma-
nently fixed. It is equally clear that if the rest of the
leaves should follow this first at uniform distances, from
the turns of the spiral being closer towards the interior,
the divergence between the successive leaves would in-
crease in proportion as they were situated more internally,
and would first acquire the dimension normal for the
species at the transit of these leaves over on to the heliacal
line. On the other side it is likewise evident that, with-
out any alteration of their divergence, the leaves would
seem to traverse a spiral, if each of them proceeded to-
wards the periphery upon the radius on which it stands
at its first origin ; for during the expansion of the axis of
the bud, its most external part extends outwards and
upwards to become the surface of the stem, the outer
end of the spiral running upon this part of the axis,
passes, as a continuation of the helix, on to the outer
surface of the stem, and the succeeding leaf advances just
so much nearer to the end of the spiral line, not because
it makes a lateral movement upon it, but because the
spiral line is abbreviated in the direction towards the leaf,
and its point of transition into the helix advances nearer
to the point at which the radius, on which the leaf stands,
intersects the circumference of the stem. Which of these
processes occurs, whether the leaf actually advances late-
rally, or the motion is only apparent, may be decided by
investigation whether or not the divergence of the leaves
alters. Now I believe that it will be found on examina-
tion of the position of the leaves, that they exhibit the
same divergence in the terminal bud as on the stem, and
that the arrangement of the leaves passes with uniform
divergence from the helix into the spiral, and is continued
in this ; while, according to Meneghini's idea, the diver-
gence must increase in the spiral. The distance from
a leaf to its successor on the spiral will naturally be

64 THE PALM-STEM:

shorter, looking at its absolute size, the nearer it is to the
inner end, but the angular divergence of the leaves will
be the same. If this be the case, and I believe that all
investigations hitherto made testify that such is the con-
dition, it is evident that the movement of every leaf takes
place in the radial direction, and is simply a consequence
of the elongation and expansion of the axis ; that the
motion of any leaf in a spiral direction is only apparent,
and results from the fact that the spiral line has no defi-
nite place upon the upper surface of the bud, but its
point of origin continually advances in the direction in
which the leaf-spiral runs, into the circle which is formed
by the connexion of the cylindrical surface of the stem
with the depressed surface of the bud. If such be the
condition, the leaves must appear to traverse the outer
turns of the spiral more rapidly than the inner, since the
same angles of divergence correspond to larger segments
of turns in the outer parts of the spiral, while, according
to Meneghini's theory, the contrary occurs : the motion of
the leaves diminishes in regard to the angle of divergence
in the outer turns of the spiral, but in regard to the spaces
passed through remains the same, and thus must appear
to become slower. Since, although I had investigated
the terminal buds of some large Monocotyledonous plants
in reference to this point, and had found no notable
deviation with regard to the divergence of their leaves,
my judgment on this matter could not be nearly so va-
luable as that of my honoured friend, Professor Alexander
Braun, I wrote to ask him whether he had ever met with
cases of such a deviation in the divergence in the terminal
bud as is required by Meneghini's theory, and obtained
the answer, that he is indeed of opinion that it is very
difficult to settle this point with certainty by direct obser-
vation, but from his own researches, he believes the diver-
gence in the terminal bud to be the same as in the stem.
From this important confirmation of my views, it appears
to me that the entire doctrine of an actual spiral motion
of the leaves, and the deduction from it of the oblique

APPENDIX. 65

direction of the fibres, must be thrown aside. There are
still further reasons against Meneghini's asserted lateral
motion of the leaves. If the oblique direction of the
fibres were caused by such a motion, it is evident that all
the fibres ought to run in the same direction (right or
left), since this movement of the leaves must have the
same effect as a twisting of the stem. Therefore, if a
Palm-stem be split longitudinally, the split surface ought to
follow that spiral in which the fibres run in the stem, and
no fibres should be torn by such a splitting, as they would
all be homodromously curved in their course downwards
from the centre of the stem. I investigated this condi-
tion in a stem of the Brazilian Palm, which has lately
been imported for manufacturing purposes, in lengths of
about seven inches. The split surfaces of this stem never
run in an oblique direction, but always parallel with the
axis, and a considerable resistance is offered in the process
of splitting, since, in the split, not only are fibres running
parallel separated, but a very great number of the fibres
of the stem must be torn across, because one portion of
the fibres runs obliquely through the stem from right to
left, and another in the opposite direction. Thus, in
reference to this oblique direction of the fibres, exactly
the same condition occurs in the Palms as in Dractena,
Yucca, and still more evidently in Xanthorrkoeq, in which
the fibres running right and left lie in alternating layers
surrounding the stem, exhibiting in the cross section some
resemblance to the annual rings of a Dicotyledonous stem.

It is sufficiently shown by the preceding observations,
that the explanation given by Meneghini of the oblique
course of the vascular bundles cannot be correct, either
in Dracaena, or in the stems which have aniplexicaul
leaves ; we must seek the reason of it, not in mechanical
causes, but in the organic activity of the plant.

In reference to the vascular bundles of the root,
Meneghini observed that in a young Chamcerops they
passed directly into the vascular bundles of the leaves ;
but that in the roots breaking through higher up on an

60 THE PALM -STEM :

old stein, the condition was different, the greater number
of their vascular bundles being lost in the outer woody
layers of the stem, only solitary fibres curving upwards
or downwards, to run further beneath the rind. In the
roots of many other arborescent Monocotyledons he con-
stantly saw the vascular bundles, if the root were still
young, spread out like a star over the woody layer of the
stem; in older roots, on the contrary, they penetrated
deeper into the stem, where they then ramified and lost
themselves. Meneghini derives these differences from
the different conditions of the course of the sap ; the
main root (tap-root) and the lowest roots of the stem are
formed at the same time, and with the help of the same
currents of nutrient sap, as the vascular bundles of the
stem, the latter, therefore, pass directly into the fibres of
the former ; but when roots are formed on the older
parts of the stem, the currents of sap cause a stellate
expansion of their vascular bundles on the surface of the
woody mass of the stem.

Unger has given some contributions to the knowledge
of the Monocotyledonous stem, having, in his researches
on the Dicotyledonous stem (Ueber den Ban und das
Wachsthum des Dicotyledonenstammes, 1840, p. 35),
taken a comparative glance at the structure of the Mono-
cotyledons, and, more particularly, subjected the stem of
the Aloinea (Draccena, Aletris, Yucca, Agave] to a more
minute investigation. With regard to the course and
anatomical peculiarities of the vascular bundles, Unger
agrees in general with the description I have given, and
thus nothing need be said on this point. But from his
investigation of the course of development of the vascular
bundles, he has been led to propound a definite opinion
as to the import of their proper vessels, while I had let
this point remain untouched. He traced the develop-
ment of the vascular bundles principally in the Aloinece,
and states that their rudiments are uniformly cellular, that
the other systems (the wood and liber portions) make
their appearance subsequently at the inner and outer

APPENDIX. 67

sides, and that their intermediate portion is to be regarded
as their most essential part, since it is not merely that
first originating, but also, in all stages of metamor-
phosis of the vascular bundle, persists under the same
form of elongated cells, resembling milk-vessels.

In reference to the blending of the vascular bundles
with each other, Unger distinguishes the proper blending
(coalifus), which consists in the actual fusion into one
vascular bundle, from the mere apposition (symphysis) ,
in which the vascular bundles are attached together by a
dense parenchyma. According to Unger, true blending
does not occur in the Aloinece (with the exception of
Yucca gloriosa and Agave Americana), while in other
families, especially in the Scitaminese and Bromeliaceae,
it is frequent.

Of the systems of vascular bundles of the buds and
roots of the Grasses, it is stated, that they are special to
these organs, and do not pass into them from the stem ;
this is particularly evident in the roots, but in the vascu-
lar bundles of the buds also, a mere anastomosis with
the vascular bundles of the stem occurs, and only indivi-
dual ones become mingled with those of the stem.

With regard to the period at which the vascular
bundles running to a leaf originate, Unger found that the
slender fibres occurring in the Palms and Grasses, which
do not enter into the interior of the stem (at least, cer-
tainly not in the Grasses), are of later origin than the
stronger vascular bundles which run from the inner layers
of the stem into the leaves.

Lestiboudois expresses an opinion very different from
that of his predecessors, in reference to the mutual blend-
ing of the vascular bundles, in his very^aluable treatise
on the Structure of the Stems of Plants (Etudes sur 1' Ana-
tomic et la Physiologic des Vegetaux, 1840), in which he
has wholly relinquished his earlier views of the structure
of the Monocotyledonous stem.

The Palm-stems examined by Lestiboudois belonged
to species, the systematic names of which are unknown

68 THE PALM-STEM :

to him ; they were chiefly two stems, imported for com-
mercial purposes, one of which has black, the other red
fibres. Lestiboudois believes the slender fibres of the
outer layers to originate partly from the cellular rind, and
partly in the form of branches of the larger fibres ; and
he states that, in consequence of manifold ramifications
and lateral connexions, they form a continuous network.
In the upper end of the fibres, entering the leaf, he like-
wise finds not only, frequently, a division of them into
several branches, entering separately into the leaf, but
also when they enter undivided into the leaf, dense rami-
fications which become blended with other fibres. He
further assumes that the fibres are not all formed at the
periphery of the stem, but that from the central vascular
bundles, and from those running in the hard, woody
layers, slender fibres, like those lying beneath the rind,
run out ; and from this he draws the conclusion that it is
clear that all fibres are destined to produce new fibres.
The particular fibres do not, indeed, run in unbroken
continuity through the whole stem, but the fibres of the
different parts of the stem are so connected that they
spring out from one another, and before they emerge into
the leaf, give off branches, which are destined to replace,
in the upper part of the stem, the fibres emerging into
the leaves lower down. From this it is clear, that in the
Monocotyledons the life is not exclusively seated in the
outer layers ; if the stem be cut into all round, down to
the hard layer, in a Yucca or Aloe fruticosa, the plants
will live, unhurt, for many years ; the only phenomenon
which appears in the wound, being the formation of a
protruding collar on the upper border of the incision,
from out of which numerous roots frequently break forth.
Before I follow the author further in his exposition, I
have to state, that I am very far from implying a doubt
of the correctness of his observations, but yet I cannot
forbear from questioning the universality and frequency
of these ramifications in the vascular bundles of the Palms.
I have stated above, 'that in particular stems of the Cocos-

APPENDIX. 69

like division, for instance, in Lepicodaryum gracile, I had
found many slender evascular fibres among the perfect
vascular bundles, in the interior of the stem ; in the stems
which I examined I could not trace the origin of these
fibres, and therefore cannot say whether they were formed
by ramification of the vascular bundles, as in Lestiboudois'
stems; in any case, however, the occurrence of these
fibres is quite unusual, and to be regarded as an anomaly
in the stems of Palms, and I found none of them in the
Palm-stems which are met with in the shops with us, and
which, doubtless, belong to the same species as those
which Lestiboudois calls "the Palms with black fibres."
That the vascular bundles do not all end below in a slender
fibre lying beneath the rind, but that it does happen a
fibre becomes blended with another at its lower extre-
mity i. e. in tracing the fibre from below upwards, it
appears like a branch of another fibre I have likewise
mentioned; but this condition occurs only in very few
fibres. As to the condition of the multitude of thin,
lower fibres lying beneath the rind, whether they are lost
in the cellular tissue or blended with other fibres, I am
unable to say, for the solidity of the tissue in this situation
rendered it impossible for me to make out this point ac-
curately. An anastomosis of these ends of the fibres into
a connected reticulation (as in Yucca, Xanthorrhcea), de-
cidedly, I believe, does not occur ; only with the existence
of such would such a universal connexion of the fibres
with each other occur, as Lestiboudois claims for the
Palms. I cannot find in the Palms a connexion and
ramification of the upper parts of the fibres, like that
described by Lestiboudois, I therefore think that their
assumption by Lestiboudois is rather deduced from the
analogy of the Palm-stems with those of Yucca and Aloe,
than founded on actual extensive researches on the former.
Lestiboudois found this ramification of the fibres in a
high degree in Yucca, Aloe fruticosa, and especially in
Pandanus. When he deduced from these observations
the conclusion that the fibres of the Monocotyledons are

70 THE PALM -STEM :

formed in a different way from those of the Dicotyledons,
originating from ramification of the old vascular bundles,
while in the Dicotyledons the new fibres are deposited
between the bark and the ends of the older fibres which
emerge into the leaves, this conclusion is not justified
by the anatomical character of the Dicotyledonous stem,
since it leaves wholly out of sight the fact, that in very
many Dicotyledons the lower end of some of these vascular
bundles stands at least in quite as close organic connexion
with other vascular bundles as in the Monocotyledons; so
that the younger vascular bundles have much resemblance
to those of the Fern-stem in respect to the arrangement
and union.

Lestiboudois devoted especial attention to the roots of
the Monocotyledons ; from the examination of them he
drew two conclusions : 1 . Their vascular bundles are not
formed by gradual elongation of those of the stem.
2. Their growth is endogenous, for their vascular bundles
are perfected from without inward, and new vascular
bundles originate in the interior of the root.

Schleiden (Wiegmann's Archiv, 1839, p. 220; Grund-
ziige der wiss. Bot. i, p. 220) directs attention to the
point that the vascular bundles are developed in a similar
manner in the Monocotyledons and the Dicotyledons, the
part turned towards the interior being that which first
originates, and he considered it to be the only thorough
distinction between Monocotyledons and Dicotyledons,
that in the latter the formation of new elementary organs
proceeds indefinitely in the cambium-layer of the vascular
bundle (whence he calls these vascular bundles unlimited},
while in the former this development ceases at a certain
epoch (limited vascular bundles), and the elementary
organs of the cambium-layer assume the peculiar form in
which I have described them under the name of proper
vessels. With regard to this distinction, the knowledge
of the facts on which it depends is not new, only the ex-
planation, in so far that Schleiden quite correctly explains
the continuous deposition of new layers of wood in the

APPENDIX. 71

vascular bundle of the Dicotyledons as a growth belonging
to the vascular bundle itself, while I had derived it from
the interposition of the lower ends of younger vascular
bundles, running to leaves situated higher up, between
the liber and wood of the old bundles ; a theory against
which Unger, also, has declared himself with convincing
reasons (Ueber den Bau und das Wachsthum des Dico-
tylenstammes) . Perhaps it may not be superfluous to
remark, that this distinction only refers to part of the
vascular bundles of the Dicotyledons, since the part en-
tering the leaf and traversing it is likewise limited in its
increase of thickness.

I now turn to Mirbel's excellent essay on the Structure
of the Date-palm (Comptes rendus de 1' Academic des
Sciences, 12 June, 1843). In 1839 the author went to
Algiers, in order to have an opportunity of examining a
full-grown Date-palm in a fresh condition ; and as the
result of four years' labour, directed especially to the in-
vestigation of the terminal bud of this Palm, he laid
before the Paris Academy a theory of the structure and
growth of the Monocotyledons, which is opposed to my
statement in several points.

With reference to the connexion of the roots with the
stem, Mirbel agrees on the whole with my views, but
assumes a more intimate connexion of the vascular bundles
of the roots with those of the stems, than I met with in
my researches. According to his statement, the fibres
which come from the middle and neighbouring parts of
the root, penetrate into the interior of the stem, between
the vascular bundles of the latter, and lose themselves
among them, in such a way that their extremities cannot
be accurately made out, while fibres derived from the
circumference of the expansion formed by the root in the
stem, are distributed upwards and downwards, in the
superficial layers. In reference to the latter, Mirbel thinks
that they probably furnish contributions to the suckers
which break forth abundantly from the base of the stem
in the Date-palm and Chamcerops; of the former, Mirbel

72 THE PALM-STEM :

thinks they may stand in connexion with the leaves.
When Mirbel then proceeds, that my statements as to
the relations of the vascular bundles of the roots to those
of the stems are not contradicted by this, but only carried
out to their real extent, I am far from calling in question
the accuracy of the observations made by Mirbel on the
Date-palm, but must defend myself from the construction
which must almost necessarily be attached to Mirbel's
expression, that, namely, my observations, and the con-
clusions to be drawn from them, had been essentially
modified, and that a more direct transition of one part
of the vascular bundle of the root into that of the stem
occurs universally. The essential point which I established
by my observations on the roots of Monocotyledons, is
the fact that the root is an independent structure, possess-
ing its own special system of vascular bundles, that these
are not gradually elongated outwards, towards the point
of the root, but also increase in length in the posterior
extremity, lying in the stem, penetrate into the stem from
the cellular mass which constitutes the basis of the root-
bud, form an interlacement with the vascular bundles
of the stem, and become attached to them.

All doubt as to this independence is removed by the
investigation of the root-bud, for its vascular bundles are
originally entirely separate from those of the stem ; but it
is also perceptible, subsequently, that at least the greater
proportion of them are not directly continuous with the
vascular bundles of the stem, but, with an alteration of
their structure, an interlacement is formed, in which they
apply themselves to the sides of the vascular bundles of
the stem, without actually passing over into them. The
bundles of the branches of the root stand in the same re-
lation to those of the root as the latter do to those of the
stem. On account of the smaller number of vascular bun-
dles of the root-branch and the root, and the very different
diameter of the vessels of these two organs, it may be ob-
served, with the greatest certainty, that an immediate
transition of either set of vascular bundles into the other

APPENDIX. 73

does not exist. This investigation is far more difficult at
the place of union of the root with the stem ; but in a
few Palms, particularly in a Cocos stem, in which the
cellular tissue had been destroyed by rotting, I examined,
at the point of entrance of several roots, the interlacement
which their vascular bundles formed with those of the
stem, and traced separately all the vascular bundles which
ran through these, without finding a transition of a root-
bundle into a stem-bundle ; I therefore think that I may
safely deny the occurrence of such a transition, at least in
these cases. At the same time I do not wish to question,
that in other cases part of the vascular bundles entering
the stem from the root become mingled with the bundles
of the former, and run on with them, as I have myself
stated this to be the case with the fibrous bundles, which,
in some Palms, lie with their lower extremities in the
rind of the root, and as Meneghini found in the isolated
fibres of the woody portion of the roots of Chamcerops ;
but I believe I may deny that this condition occurs always
and necessarily, and that a conclusion adverse to the in-
dependence of the vascular-bundle system of the Mono-
cotyledons is to be deduced from it.

In regard to the structure of the stem, Mirbel directed
his attention chiefly to the settlement of the question,
whether the vascular bundles grow downwards from the
leaves into the stem, or are developed in the contrary
manner. He is in doubt whether, in my description of
the Palm-stem, where I have described the course of the
fibres downwards from the leaf, I merely intended to de-
note the mechanical conditions of their position, or the
direction in which they are formed. I scarcely think that
I could have expressed myself more distinctly on this
point, that I meant only the former. I was unwilling to
propound any definite opinion on the second question,
because, in the absence of sufficient material, I was unable
to solve it with certainty.

Mirbel first investigates the question, whether the vas-
cular bundles run down from the leaves to the base of the

74 THE PALM -STEM :

stem, and founds his answer on the following considera-
tions. His Date-palm was 18 '60 meters high, the base,
clothed with roots, 34 centimeters in diameter ; above the
root-bearing part the surface of the stem had suffered
from the influence of the atmosphere, and its diameter
here amounted to 25 centimeters; the upper part was
covered with leaf-scars, and almost cylindrical. Mirbel
then concludes, that if the fibres originated in the leaves,
and all reached down to the base of the stem, or, on the
other hand, if all the fibres originated at the base of the
stem, and ascended up into the leaves, the stem must
necessarily have a conical form, on account of the accu-
mulation of the fibres at its lower end. In like manner
is opposed to either of these views a circumstance, which
was undoubtedly known to me, namely, the spindle-shaped
expansion of many Palm-stems in the middle, which I
could not explain according to my theory, but which
affords nothing remarkable now he has found that the
fibres originate at all heights in the stem. The most
certain evidence, however, against the opinion that the
fibres run from the leaves to the base of the stem, is
furnished by the following exact measurements. In the
Date-stem he examined there were 337 leaf-scars on a
length of one meter, the entire stem had, therefore, borne
about 6268 leaves. At the base of one petiole Mirbel
found 500 fibres 1 millimeter thick, and 400 fibres ^th
of a millimeter, which he estimated equal to 44 of the
larger fibres ; for the vagina of the leaf he reckoned 100 ;
therefore, altogether, 644 for a leaf, and for all the leaves
of the stem added together, 4,036,592 fibres. Besides
these, account must also be taken of the fibres which run to
the spathes and flower-stalks, and, moreover, the enormous
number of capillary fibres which occupy a considerable
space in the hard and firm crust of the oldest portions of
the stem. Leaving these last out of sight, the fibres
running to the leaves furnish a sufficient proof against
my theory, since, if these fibres ran down to the base of
the stein, the cone formed by them at the bottom would

APPENDIX. 75

be 2*01 meters in diameter, and 6*33 meters in circum-
ference, while, in reality, the said stem was only 25 centi-
meters in diameter above the root -stalk.

Before I follow Mirbel's researches further, I must
subjoin a few words to the foregoing. In the explanation
of the structure of the Palm-stem I have called attention
to two circumstances, to the course of the fibres, and to
the alteration of their structure and size in different parts
of their course. To the latter point, Mirbel, in advancing
the preceding calculation in opposition to my view,
paid no attention, but it was necessarily required that
lie should have done so, to afford any useful result.
Mirbel says the Palm-stem swollen in the middle is a
complete contradiction to my theory. The fact that such
stems occur was well known to me, but by itself it was
not of sufficient importance to give rise to a different
theory. That with the development of the stem and the
increase of the force of its vegetation, the upper leaves
are larger than the lower, and a greater number of fibres
run from them into the stem, than from the lower ; that
the stem must consequently acquire greater diameter at
the base of these leaves than at the base of the lower
leaves all this is exceedingly simple. This increased
thickness must be continued to the lower part of the stem,
and the latter thus acquire a conical shape if the vascular
bundles run down without change of thickness to the
base of the stem, but by no means if, at the place where
they appear beneath the surface of the stem, they be-
come so slender, that in spite of their accumulation they
still form no equivalent for the greater size of their
upper ends, situated higher up in the stem. The plates
of my ' Anatomy of the Palms' afford evidence of how
considerable this attenuation is ; they are drawn with the
Sommering's reflector, the relative sizes of the various
parts of the same figure are therefore accurate, and a
comparison of the vascular bundles with the thin fibres
lying beneath the rind, shows that the transverse section
of the latter is frequently a hundred times smaller than

76 THE PALM-STEM :

that of the former. With this considerable attenuation
of the vascular bundles at their lower extremities, with
the more distant position of the upper parts of them in
the interior of the stem, and with the crowded position
of their lower filiform extremities, even if the fibres did
run down in the stem, the lower part of this would only
undergo a relatively slight increase of thickness, and a
spindle-shaped thickening in the middle would still always
be possible. Mirbel, on the other hand, reckons as if all
the vascular bundles arrived at the base of the stem of
the same thickness as they are when they emerge from
the leaf, and comes to the conclusion, that in his stem
the mass of these fibres would form a cylinder, which in
the transverse section would display a surface nearly sixty-
four times larger than the cross section of the stem
actually was. To make a calculation of any use, it is ne-
cessary to settle the thickness of the lower end of the
fibres. I have before me the discoid slice of a full-grown
stem of Phoenix, 34 centimeters in diameter, in which
the fibres lying beneath the rind average 0*127,
therefore nearly | of a millimeter in diameter. Conse-
quently, if we assume with Mirbel that the upper
and middle portions of the fibres are 1 millimeter in
diameter, the cross section of about 64 of these fibres
would equal it. If, then, all the vascular bundles reached
the lower end of the stem under the form of such slender
fibres, taking the number of vascular bundles of MirbeFs
stem, the sum which he obtained must be diminished
sixty-four times, i. e. the mass of these fibres would form
a cylinder of the thickness of the Palm-stem Mirbel ex-
amined. I do not at all intend to attach any weight to
this calculation, its incorrectness is too evident, since, ac-
cording to it, the stem would consist of a compact fibrous
mass; but it shows that a calculation, if not based on
much safer grounds than Mirbel's rests on, cannot lead
to any useful conclusions.

I admit unconditionally that my statement, that the
vascular bundles run down, in the form of fibres, to the

APPENDIX. 77

base of the stem, is incorrect as regards the Palms. I
was led to the assumption by the investigation of too
young specimens (for I only had such in an entire con-
dition, and the portions of full-grown Palm-stems at my
disposal were merely short pieces), as well as by too wide
an extension of the analogy with the stems of Draccena,
Yucca, &c. The grounds on which I now hold the view
that the fibres run down to the base of the full-grown
stem is false, are anatomical. It is clear, that in the
transverse section of an old Palm-stem, we ought to find
beneath the rind the fibres which are only just beginning
to be formed, as Moldenhawer states that he did, although
he had indeed no large stems to investigate ; now in the
examination of sections of old stems I have not seen
this, but all the fibres lying beneath the rind were com-
posed of thick-walled cells, consequently were old. A
subsequent growth of the fibres beneath the rind of full-
grown stems, therefore, does not take place. The matter
is altogether different in respect to the stems of Aletris,
Dracana, &c. ; here there is a layer beneath the rind
perfectly comparable with the cambium-layer of the
Dicotyledons, in which parenchymatous cells and fibrous
bundles grow subsequently, which formation of new
structures gives rise to continual increase of thickness of
these stems. It is proved by this that the fibres termi-
nate higher up in the stem in the Palms, and that in
them must exist a condition similar to that in many
Dicotyledons, where the vascular bundle can only be
traced down through a certain number of internodes.

Passing now to the most important part of Mirbel's
treatise, the examination of the terminal bud, he states
that he found two slits, one above the other, in the centre
of the flattened, concave, excavated apex of the stem,
composed of nascent cellular tissue (to which he applies
the name of pfiyllopliore], these slits dividing the cellular
tissue into superimposed layers. Of these layers each,
according to his views, represents a nascent leaf; the
upper one is elevated into a vesicle, and this becomes

78 THE PALM-STEM :

torn away, in a circular direction in the greatest part of
its circumference. The isthmus is developed into the
petiole, the upper part of the vesicle becomes erected,
acquires the shape of a spoon, and is converted into the
leaf, the vagina of the leaf appears to grow out from the
wound which the torn leaf leaves upon the phyllophore.
The leaf acquires the form of a hood, its border having
an irregular thickening ; the two lateral halves of the hood
are formed of the two series of leaflets, and the thickened
edge, which unites the points of the leaflets, subsequently
becomes absorbed.

The description of this process does not agree in the
least with what I observed in regard to the earliest period
of the formation of the Palm leaf. I examined, in reference
to these statements of Mirbel, the terminal buds of
Phoenix and Cocos flexuosa^ but found, as in other Mono-
cotyledons, such as ^ave, Yucca, no trace of origin of
the leaf under the form of a circularly torn vesicle as de-
scribed by Mirbel, but saw the leaves shoot forth from
the axis in the form of obtuse papillae. This papilla is at
first narrow, in proportion to the portion of the axis on
which it stands, since the first-formed part of it corre-
sponds to the apex of the future leaf; the further it is
developed, the more the base rises from the surface of the
stein, so that in the Palms an indication of the vagina of
the leaf is visible at a very early period. I cannot under-
stand how Mirbel came to the idea that the leaf originates
in the form of a vesicle, and that several such vesicles lie
one above another ; he must have been led to this view by
a longitudinal section which did not pass exactly through
the axis of the bud, and thus have met with sheaths of
young leaflets (which in the inner parts of the bud are not
cylindrical, but have the lower part spread out almost flat),
and have taken them for the rudiments of the whole leaves.
I deduce another reason against the assumption that the
leaflets originate from closed vesicles, from the observation
of a monstrous formation which I found in a branch of
Phcenix, the axis of which had grown out to the length

APPENDIX. 79

of some two inches. About six leaves on this had no
amplexicaul vaginas, but the lower sheath-like portion of
all the leaves formed a connected lamella, passing round
the stem in a spiral line, from the upper part of which
normally -formed leaves were given off at intervals. The
upper leaves had perfectly closed sheaths. Similar con-
fluence of several leaves into a continuous leaf-spiral have,
as is well known, been repeatedly observed in plants with
verticillate leaves, ex. gr. in Casuarina, whether they have
elsewhere occurred in amplexicaul leaves I know not ; in
any case, the case just described appears to me incon-
sistent with the idea of the origin of the leaves in the
form of closed vesicles.

The development of the originally simple leaf into a
pinnate leaf is very peculiar in the Palms. Since Mirbel
rather indicates than describes this process, a more minute
explanation of it will, though not very closely connected
with the object of this essay, perhaps be acceptable to
many readers. De Candolle, in his ' Organography '
(i, 304), has already observed, that the division of the
Palm-leaf into pinnae, or into lobes of a fan-shaped leaf,
takes place in a manner altogether peculiar, namely, by
a tearing up of the structure. De Candolle looked at this
process much too roughly ; he evidently made his ob-
servations on a leaf already developed to a considerable
degree ; I was therefore quite justified in rejecting this
notion of a mechanical description in my ' Anatomy of
the Palms/ but in like manner did not go far enough
back in the investigation of the young leaf, in its earliest
stages of development, to enable me to give a satisfactory
explanation of the process. I had indeed correctly made
out that the separation of the pinnae is completed long-
before the unfolding of the leaf, and that they are not
connected together in the bud by leaf -tissue, but by a
loose parenchyma, which is blended in a very narrow line
with the border of the leaf, is connected with the pubes-
cence of the leaf, and dries up and falls off with it,
whereby the leaflets are allowed to separate from each

80 THE PALM-STEM :

other and become free ; when, on the other hand, I ex-
plained this tissue as a peculiar form of the pubescence,
I was wrong, as the following observations will show.
I traced the development of the leaf in Pkcenisc (pi. I,
figs. 9-13) and Co cos fleocuosa (figs. 1-8) ; in both, the
young leaflets (figs. 1, 12, 13), until they attain the
length of about five millimeters, are composed of a con-
nected tissue, which in the middle, as rudiment of the
future petiole, is thicker, and runs out into a relatively
thin border at each side. At a later period a smooth
furrow is formed between the thickened mid-rib and the
margin of the leaf (fig. 2), at the bottom of which are
subsequently met with, nearly approached, somewhat
excavated cross-striae (figs. 3, 4), the tissue of the leaf,
however, being still continuous. These cross-striae are
afterwards converted into narrow slits (figs. 5-11), which
in Cocos flexuosa penetrate the entire thickness of the
leaf, so that they are visible on the upper and lower sur-
faces (fig. 7 o). The further development shows that the
part lying between each pair of slits becomes perfected
into a leaflet, and in a cross section (fig. 7 c], or, still
better, in a longitudinal section, it is perceived that these
pinnae are folded together, and that the mid-rib, at which
the fold takes place, is in Cocos on the upper surface, so
that consequently twice as many slits are visible at the
under side of the leaf as at the upper (fig. 7 c). The
margin of the leaf, at which the points of all the pinnae
are blended, forms a continuous cellular mass, which ends
externally (fig. 8*) in an acute angle (the border of the
previously undivided leaf). With the advance of the
development of the leaf this cellular mass dries up, and
is thrown off in the form of a brown filament, by which
the leaflets are set free. In Phoenix (figs. 9-13) the
matter is rather different, as the mid-rib of the leaflets is
turned toward the under side of the leaf, and the cellular
mass which connects the pinnae is not merely blended
with their summits, but is continued over the whole of
the upper surface of the leaf as a rather thick membrane,

APPENDIX. 81

and is blended with the upturned margins of the pinnae,
whence the slits between the latter are only visible at the
under side of the leaf. The leaf, therefore, originates as
a continuous mass, and the pinnae owe their origin to an
actual division of the leaf : the division, however, does
not advance from the margin of the leaf toward the mid-
rib, but relates only to the surface, not affecting the
border, nor, in Phcenicc, the upper layer of the tissue of
the leaf. This permanently undivided mass of cells is
distinguished from a true pubescence, to which it bears
much resemblance, by its origin, since it is not a growth
from the surface of the organ, but forms an actual part
of the tissue of the leaf, as well as by the circumstance
that in some of the Palms, for example in Phoenix (but
not in Cocos), vascular bundles run into it.

Return we from this digression, to Mirbel's description
of the bud. He states that there run through the tissue
of the bud a countless number of transparent, very deli-
cate fibres, which converge from the whole internal
periphery of the stem toward the central portion of the
phyllophore, where their upper extremities approach the
young leaflets, sooner or later to enter into direct con-
nexion with them. In a few instances he has hit upon
these fibres at the moment when they were running into
the weakly -indicated rudiments of the leaves. Those
physiologists who believe that the fibres run down from
the leaves had certainly never an opportunity of seeing
the terminal bud of a vigorous Date-palm, or they would
have left nothing for him to do. A glance is sufficient
to convince that the upper ends of these fibres are very
young compared with the lower, that they consequently
grow from below upwards. If they sprang from the
leaves, they must be old and hardened at their point of
origin, long before they reached the base of the stem.

This proposition contains the nucleus of Mirbel's doc-
trine of the structure of the Monocotyledons. The remark
that earlier labourers at the anatomy of Palms had no
terminal bud of a vigorous stem to examine, is unfortu-

82 THE PALM-STEM :

nately, in my own case, only too well founded, and if I
venture, notwithstanding that I have not the appliances
of the Paris Academician, to subject his researches to a
criticism, I know well the difficulty of my undertaking,
yet I hope that the reasons which I have to adduce will
not be without weight. That the first glance at the sec-
tion of a Palm -bud will not force the conviction upon every
one, that the fibres grow from below upwards, is proved
most decisively by Gardner saying, that the greatest
sceptic would only need to see a longitudinal section of a
Palm-stem bearing its leaves, to be convinced that its
woody substance is formed from the leaves (Annals of
Nat. History, vi, 61.) I unfortunately had no larger
buds at my disposal than the terminal buds of a stem of
Cocos fleocuosa about two inches thick, and of young
stems of Phoenix. From an examination of these buds 5
made in reference to Mirbel's views, I have nothing to ob-
ject to the correctness of his anatomical statement, that in
Phoenix the vascular bundles running to the young leaves
of the bud, are harder and more developed below the phyllo-
phore than in it, where they are in a soft, gelatinous con-
dition. But does the conclusion deduced from this fact by
Mirbel, that the lower parts of the bundles are older than
the upper, necessarily follow from it ? At first sight,
undoubtedly, since it is a general fact that young woody
bundles are soft, old ones hard. But it is a totally different
question, whether the hardness of them is connected simply
with their age, or at the same time and in a higher degree
with the stage of development of the part in which the
vascular bundles lie. Mirbel assumes the former ; I be-
lieve the latter may be proved. Calling to mind that in
the articulated plants with elongated internodes, as in
the Grasses, Pinks, in Ephedra, a yet imperfectly-deve-
loped internode is already completely hardened above,
where it is exposed to the air, while below, so far as it is
inclosed by the sheath of the lower leaf or verticil of leaves,
its internode is still of an herbaceous softness, we have
here a case which stands in the most glaring contradic-

APPENDIX. 83

tion to the fundamental proposition on which Mirbel rests.
Mirbel says, because in the Palms the vascular bundles
are softer above than below, their lower portion is older,
they consequently grow from below upwards ; with equal
right he would assume the reverse if he split an internode
of Zea Mays. Moreover, if we remove the leaves from a
terminal bud of Phoenix, we find in its half-developed
leaves, which have attained a length of about 1 to 3 lines,
the part of the petiole projecting from the bud already
green and firmly lignified, while the base of the petiole
and sheath are uncoloured and very soft ; the vascular
bundles of the upper part are found completely lignified,
those of the lower portion still half gelatinous and trans-
lucent, in short, we find here between the upper and
lower ends of the vascular bundles (only in reversed order)
the same differences as in the vascular bundles of the stem.
Shall we conclude from this that the vascular bundles of
the leaves grow from above downwards, that the upper
part of them is so much older than the lower ? This no
one will wish to assert, since in the Palm-leaf, it must be
concluded, from the fact that leaflets still very small have
a sheath, that the growth of the leaf depends on an ex-
pansion of the very young leaf in all directions, and not
on a subsequent after-growth of its lower portion. From
these circumstances, it certainly follows that the greatest
differences of solidity and completion of structure occur
simultaneously in different parts of the same vascular
bundle, and may be caused by the transition from the
rudimentary to the lignified condition proceeding with
different rapidity in different parts of it, and going on
parallel with the unequal growth of the organs in which
the vascular bundle lies, since, in general, the upper part
of the leaf attains its full development first, and, in the
slowly-growing Palm-leaf, a long time before the lower
portion ; the same holds good in the vascular bundles of
its petiole, and the direction in which their lignification
proceeds, can by no means be regarded as an index of the
direction in which their rudiments made their appear-

84 THE PALM-STEM :

ance in their original formation. A vascular bundle
which runs from the summit of the petiole of such a half-
developed leaf, through the petiole and bud, to the outer
surface of a part of the stem situated far lower down,
may, according to what has just been said, exhibit its
complete development and solidity at both ends, which
lie in organs already completed and solidified, while its
middle portion, in its course through the lower part of
the petiole and through the upper soft portion of the stem
(the phyllophore) concealed in the bud, is gelatinously
soft, and, in reference to the anatomical completion of its
individual elementary organs, still in a young condition.
In the same proportion as these soft parts advance in
development, as the part of the phyllophore through which
the vascular bundle runs becomes part of the stem, and
the petiole and leaf-sheath attain their full size and be-
come solidified, will the soft part of the vascular bundle
also increase in size and internal completeness, till its
growth is finally terminated, and it is solidified. This,
and nothing further, do we learn from a view of the
section of a bud, and when Mirbel finds in the increase
of solidity which the vascular bundles exhibit from above
downwards, a proof of the direction in which they origi-
nate, the validity of this conclusion must be altogether
contested.

The question, in which direction the vascular bundles
are formed at their first origin, can only be decided by
direct observation of the process of their origination ;
therefore, only by microscopic investigation. Mirbel did
not neglect researches of this kind, and he states, that he
sometimes caught the vascular bundles at the moment
when they were betaking themselves into the rudimentary
indications of the leaves. In the examination of the ter-
minal buds of various Monocotyledons, especially in Iris,
Acorus Calamus, of the bulbs of Narcissus poeticus, I came
to the result, that the first indications of the vascular
bundles, in the form of transparent streaks, are to be
met with beneath the youngest leaves, in the axial portion

APPENDIX. 85

of the bud, before they appear in the leaves, and that,
subsequently, the origin of the vessels in these rudimentary
vascular bundles follows in the same direction. This
result has been fully confirmed by the researches of
Schleiden (Grundziige, ii, 189), Meneghini (Intorno alia
struttura del tronco delle Monocotiledoni, in Miscellanee
di Chimica, Fisica e Storia naturale, 1843), and Naudin
(Ann. des Sc. nat., 1844, i, 162); and after these ac-
cordant observations, there can be no doubt that the
upper end of the vascular bundle grows from below up-
ward, and that we have to seek its origin in the stem,
and not in the leaf. But it is quite a different question,
whether this process of development occurs throughout
the whole of the vascular bundle, or if its lower end,
running downward in the stem, grows in the opposite
direction. Mirbel assumes the former, and assures us that
in the Palms, the same vascular bundle has already, at its
lower end springing from the interior of the periphery of
the stem, the characters of developed wood, and exhibits
in its intermediate parts, the half-solidified condition of
the alburnum, while at its upper extremity it consists of
nascent tissue. Has Mirbel really observed this ? I here
take the liberty to doubt. According to my own observa-
tions, the vascular bundle of a Palm, the lower end of
which already exhibits a ligneous solidity, goes, not to the
rudiment of a leaf, but to one already tolerably advanced
in development ; therefore, if the portion of it lying in
the upper part of the stem be still very soft, from the
causes explained above, this can give no further result
as to the mode of its first production. We can only ar-
rive safely at such by tracing downwards one of these
vascular bundles, the upper part of which has not yet
reached a leaf, and observing its further gradual develop-
ment. From the great number and the entangled con-
dition of the vascular bundles, I was unable to do this in
a Palm-stem, and my efforts to make out this point by
direct observation totally failed. When, in spite of this,
I venture to discuss the question, and to deduce a decision

86 THE PALM-STEM :

of it from more distant phenomena, I am fully aware
that this proceeding can claim no certainty, but at most
a certain degree of probability ; I nevertheless hope that
such a mode of considering it will not be altogether use-
less. In the first place it has to be inquired, whether a
developing vascular bundle always grows in one direction,
or whether cases do not occur where its two ends become
elongated in opposite directions. In my opinion, the
latter case undoubtedly occurs, especially at the point of
insertion of a root upon a Monocotyledonous stem, and
of a branch of one of these roots upon the main trunk.
In both these cases, and especially distinctly in the latter,
we see vascular bundles originate in the cellular node in
which the formation of the root begins, the end of which
toward the point of the root, increases in length with the
further growth of the root, while the other end penetrates,
in the opposite direction, into the stem or primary root.
In an analogous manner, as is again to be observed more
clearly in the Monocotyledons than in the Dicotyledons,
the bud which is developing into a leafy branch, also
possesses its proper system of vascular bundles, inde-
pendent of that of the stem, the lower extremities of which
pass into the stem, and spread themselves out over a lesser
or greater portion of its ligneous mass. Now, these vas-
cular bundles are nothing else than the inferior extremi-
ties of the vascular bundles of this branch, and the most
ready supposition, on seeing these fibres passing over into
the stem, is that they have been developed from above
downwards. To this explanation, which immediately pre-
sents itself, it may certainly be objected, that the fact of
the passage of that vascular bundle into the stem, does
suffice to prove that this is perfected in the direction from
above downward, for the formation of that vascular bundle
in the stem itself may be caused by the presence of the
branch, through the attraction it exerts on the sap-bearing
substance of the stem, and it may grow from the stem
upward into the branch. But the direction which the
fibres lying beneath the rind of Monocotyledonous stems

APPENDIX. 87

assume when the stem is wounded, appears to afford an
evidence that their formation actually does take place
from above downward. I have before me the stem of a
Yucca, in which many branches were sawn off from the
surface during the life of the plant, and which had other-
wise undergone injuries penetrating as far as the fibrous
layer. On these wounded places over-growths have been
formed, as in a Dicotyledonous stem, in which the fibres
run down from above to the injured part, till they reach
the upper border of the wound, then deviate to each side,
run down along the lateral borders, and at some distance
below the injured spot again approach together from each
side, in a very acute angle. In this way an over-growth
is formed above and at the sides of the wound, but is
absent at the lower border. If the fibres grew upward
from below, as Mirbel assumed, they should reach the
lower margin of the wound, deviate to the side of it, and
gradually approach together again above it ; the over-
growth ought, therefore, to originate at the lower and not
the upper side of the wound. It is to be observed .here,
that this over-growth is not effected by an increasing
thickening of old vascular bundles, existing at the time of
the injury, as in the Dicotyledons, but is formed by newly-
developed bundles, which are entirely separate from those
subjacent (as in general, also, in the normal growth of
the stem, the superimposed fibrous layers are not to be
compared with the annual rings of the Dicotyledons, but
to be regarded as entirely isolated structures), which is
most clearly shown in the direction of the spiral lines in
which the fibres run, since these spirals alternate to the
right and left, in the successive layers, in a manner ana-
logous to what is found in Xanthorrhoea.

The analogy between the structure of the stem of a
Yucca and a Palm is so great, that we are justified in
drawing a conclusion from the phenomena we observe in
the former, as to the processes occurring in the latter'.
The distinction between the two stems lies chiefly in
this, that in Yucca the lower extremities of the vascular

88 THE PALM-STEM :

bundles run down to the base of the stem in the form of
a close fibrous network, and that in consequence of the
continued deposition of new fibrous layers, the stem ex-
hibits an uninterrupted growth in thickness, while in the
Palms the lower fibrous extremities of the vascular
bundles, as a general rule, remain simple, and do not run
downwards to the base of the stem. The case observed
by me in Co cos, of the solution of the vascular bundles
into a number of slender fibres (p. 9) is to be regarded
as an approximation, in the Palms, to the structure of this
fibrous reticulation. These differences occurring between
the stem of Yucca and that of the Palms are undoubtedly
not of sufficient importance to allow of our supposing an
essential difference to exist in the mode of development
of the vascular bundles. Since, then, in every Monoco-
tyledonous stem vascular bundles appear, in consequence
of the development of a branch, which, without the for-
mation of that branch, would not have originated, since
these vascular bundles form a greater mass, and spread
so much more widely over the stem from the base of the
branch, the older this latter becomes, since in Yucca the
course of the vascular bundles exhibits such mechanical
conditions in the vicinity of a wound on the stem, as
must result from a downward growth of the vascular
bundles on the stem, it is quite justifiable for me to de-
clare that Mirbel's view, that the vascular bundles of the
Palms grow from below upwards, is an opinion opposed
to the phenomena of the growth of the Monocotyledons ;
and to presume, on the contrary, that the lower portion
of these bundles is developed in the direction from above
downwards.

Meneghini came to the same result through a series of
deductions altogether different from mine. He had already-
stated, in his first paper on the structure of Monocotyledons
(Richerche sulla Struttura del Caule nelle Piante Monoco-
tiledoni, 1836, p. 77), that the formation of vascular
bundles was caused by definite currents of nutrient sap,
without, however, carrying out this view more minutely.

APPENDIX. 89

In a more recent treatise (Intorno alia struttura del
tronco delle Monocotiledoni), he sets up the following
theory, as to the connexion between the currents of sap
and the origin of the vascular bundles, and as to the
dependence of the direction in which the fibres are
developed on the course of the sap. The nascent leaf in
the centre of the bud forms the focus of the sap-currents,
which converge from the periphery of the bud to the leaf,
and give rise to the production of vascular bundles. The
portion of the vascular bundle which has originated in
this way, forms, when the leaf, in the course of its further
development, has passed out to the periphery of this stem,
the inferior portion of its vascular bundle, since the
upper part of it grows after that first portion, gradually,
and in proportion as the leaf is developed and assumes
its subsequent position. During the perfecting of the
leaf, the sap-currents running in toward it, and with
them the organization of the vascular bundles in the
cambium-layer of the stem, undergo a continually in-
creasing expansion in the direction from above downward.
When the leaf has emerged from the bud, and has
become green, the superabundant assimilated juices
flowing toward it, together with the ascending sap, flow
back, and give rise to the formation of liber-cells. The
ascending sap, which is so much the less elaborated the
nearer it is to the root which absorbs it, must influence
the metamorphosis of the cells through which it flows, so
much the more powerfully, the nearer it comes to the
point by which it is especially attracted ; therefore the
formation of the vessels, which depends upon it, begins
in the leaf and descends from here to the root. The
descending sap, on the other hand, which is consumed in
the nutrition of the cells, must lose its activity in propor-
tion as it becomes removed from the leaves. The forma-
tion of the vessels depends on the ascending, that of the
liber-cells upon the descending sap; the first, consequently,
predominates in the upper parts of the vascular bundles,
the other in their lower portions, and on double grounds

90 THE PALM-STEM :

the vascular bundles must, in respect to the organic for-
mation, be regarded as descending from the nascent leaf
in the centre of the bud.

Returning to Mirbel's exposition, the manner in which
the vascular bundles terminate below in Phwnix is not
minutely described by him. Neither have I, as already
observed, been fortunate enough to make out this point
with certainty. It is indeed found, as I stated in my
description of the Palm-stem, that a part of the vascular
bundles becomes blended with others below, or as phy-
totomists who ascribe an ascending course to them ex-
press it, are branches from other vascular bundles entering
into leaves situated lower down ; but this condition can
only be demonstrated in a very small portion of them,
and, in particular, not in the capillary fibres, in which
the vascular bundles terminate in the outer layers of the
stem. It is probable, therefore, that the majority of the
vascular bundles terminate blindly in the cellular tissue
beneath the rind. This assumption may appear to create
a difficulty in regard to the explanation of the manner in
which the sap ascends, but we find exactly the same
condition in the vascular bundles of the corona of many
Dicotyledons for example, in Laurus nobilis, Quercus,
Rosa; where the vascular bundles running into one leaf run
down the stem without entering into any connexion with
those of another leaf, and become gradually attenuated
under the form of thin fibres, which cannot be traced
farther down. In these plants, the ascending sap, when
it flows through one vascular bundle, must, to reach a
second, emerge laterally into the cellular tissue, and from
this flow into the neighbouring vascular bundles. The
condition, however, as I have already several times re-
marked, does not occur in all the arborescent Monocoty-
ledons, since in Dractena, Yucca, Xanthorrhcea, the lower
extremities of the vascular bundles grow together at their
sides, and in this way form a connected fibrous reticula-
tion over the whole stem.

Mirbel distinguishes different kinds of vascular bundles

APPENDIX, 91

in the stem of the Date-palm : 1st, those which occur in
the interior of the stem, and form the principal mass of
its wood ; 2d, capillary fibres, which lie in great numbers
in the peripherical region of the stem and the petiole,
do not occur in the interior of the stem, and are thirty-
six times smaller in diameter than the first. These thinner
fibres contain no vessels, but are composed merely of
elongated, thick-walled cells. In my description of the
Palm-stem, I have likewise directed attention to these
differences, and mentioned that the second class of fibres
are the inferior extremities of bundles, which, without
previously entering into the interior of the stem, run in
the outer layers of the stem, and are here mostly con-
verted into true vascular bundles, linger was the first
to remark that these fibres are of later origin than the
vascular bundles lying in the interior of the stem ; but
Meneghini called especial attention to the circumstance,
that, as a general rule, the different vascular bundles of
the same leaf do not all penetrate to the same depth in
the stem, and that this difference depends on the circum-
stance that the leaf, during its development, passes out from
the centre of the axis to the periphery, and each vascular
bundle only reaches that point in its internal curve, at which
the leaf was actually situated at the moment when the
organization of the vascular bundle commenced (Intorno
alia Struttura del Tronco delle Monocotiledoni, p. 12).

Mirbel brings a portion of the vascular bundles promi-
nently forward under the name of precursors (precurseurs).
He applies this name to them because they are the first
which become connected with the leaves. According to
him, their number equals that of the leaves in each step of
the leaf-spiral (de chaque pas d'helice), and they appear at
intervals, which are measured by the length of the inter-
nodes. They lie in a bundle in the centre of the stem ;
each of them proceeds singly out of this central bundle,
and passes obliquely upward into a leaf. On its way, a
number of other vascular bundles attach themselves to
it. At the point where the precursor leaves the vertical

92 THE PALM-STEM.

direction to repair to a leaf, it sends off usually one, more
rarely two or three, branches, which ascend vertically
upward, and probably run to leaves situated higher up.
This is the only example of ramification of a fibre which
Mirbel met with in the Date-palm.

The preceding, Mirbel continues, is not at all in con-
tradiction to my statements ; but this is not the case with
the circumstance that the lower ends of the precursors
do not run in a vertical direction down the stem, but
repair to the side of the stem opposite to the leaf, whence
it results that all the precursors which pass to each one
of the turns of the leaf-spiral, cross in the central bundle,
and form two cones connected at their apices one erect,
the other reversed. Whether the direction in which these
fibres diverge from the straight line is the same in all,
Mirbel does not express ; nor does he make any mention
of the fact, that Meneghini devoted especial attention to
this point, or of the explanation of it given by that author.
From his very laconic treatment of this point, which he
speaks of as the most important difference between our
works, Mirbel gives me no opportunity of making an
accurate acquaintance with and discussing his views ; for
I must confess that it is not clear to me, from his treatise,
how and why he distinguishes the fibres he calls precur-
sors from the other developed vessels, and that I know
nothing of them.

NUCLEI, FORMATION, AND GROWTH

VEGETABLE CELLS.

BY CARL NAGELI.

PART II.

TUANSLATKD FROM

SCHLEIDEN U. NAGELl's ZEITSCHR1FT P. WISS. BOTANIK, 1846.
BY ARTHUR HENFREY, F.L.S.

VEGETABLE CELLS.

PART II.

IN the First Part of this essay I endeavoured to demon-
strate, in the first place, that nuclei occur in all vegetable
cells, and that they are utricles ; secondly, that in one
kind of cell-formation (which is called parietal cell-forma-
tion), the whole contents of the parent-cell become divided
into two or more portions, and that around each of these
a perfect membrane is formed by the secretion of gela-
tinous substance, this membrane being in contact in part
with the wall of the parent-cell, in part with that of its
fellow secondary-cell. I shall hereafter find an oppor-
tunity of defending my theories against some objections
which have since been made to them. I have delayed
the continuation of the essay, which is principally to treat
of free cell-formation, because my observations have never
appeared sufficient to afford a quite positive conclusion.
Even now, in spite of all my efforts, I have only arrived
at probabilities in my results on many points, which,
however, I will not any longer keep back, but leave the
further development and establishment of the doctrine to
a happier time.

IV. FREE CELL-FORMATION.

a. Without visible nucleus.

In free cell-formation without a visible nucleus, the new
cells originate, in the contents of the parent-cells, as mi-
nute globular bodies, in which, so soon as they have

96 VEGETABLE CELLS.

acquired sufficient size, we recognise an enveloping mem-
brane and inclosed contents.

In Chlorococcus, Grev., and Htemato coccus, Ag., the new
cells appear as green (in the former genus) or red (in the
latter) globules, in which we only perceive a definite out-
line. When they leave the parent-cell and become larger,
we distinguish a colourless membrane inclosing coloured
contents. I have never been able to detect any sign of a
nucleus. It appears as if merely small isolated portions
of the contents became agglomerated, assumed a perfectly
spherical form, and produced a membranous coat. The
rest of the contents of the parent-cell are gradually dis-
solved.

Valonia utricularis, Ag., consists of one large cell. On
the inner surface of its membrane lies the mucilaginous
layer. The germ-cells (Keimzellen) originate in this. At
first they have the appearance of minute drops of mucilage.
The membrane cannot be detected until they have become
larger, granular, and green.*

The origin of the germ-cells (spores) in the thecse of
Fungi and Lichens is accompanied by the same phe-
nomena. At first they present themselves, sometimes in
the form of homogeneous drops of mucilage, sometimes
as little hollow globules, the former in clear, diluted, the
latter in dense, mucilaginous, cell-contents of the theca.
In both cases the nascent spore-cell originally exhibits
only a definite outline, and a distinct membrane only at
a subsequent epoch, f No nucleus is visible in these ; at
a later period, when the cell has increased in size, and
the contents have become transformed, a nucleus is seen
in some instances.

In Coleochcete scutata, Breb., the germ-cells appear in
the parent-cells, at first as little green, homogeneous glo-
bules, on which a membrane afterwards becomes visible.
A nucleus may sometimes be detected in the fully-de-

* For a more minute account, see Nageli's New System of Algae. (Die
Neuen Algen-systeme, &c. Zurich, 1847.)
f See Nageli, Linnrea, 1842, p. 257, PL ix, figs. 32, 34, 41, 45.

CKLL-FORMATION. 97

veloped cell in germination. The examples just given of
free cell-formation, without visible nucleus., agree with each
other in the point that the new cells first appear as little
spherical, homogeneous portions of the contents of the
parent-cell, around which no membrane, but merely a
definite outline, can be perceived. They are, like the
contents of the parent-cell, coloured (green or red) or
colourless. Not until they acquire greater size, and in
most cases, only at the time when they become granular,
does the membrane become distinctly visible. A nucleus
is never seen in the earlier stages, and appears sub-
sequently only in isolated cases.

I have made a second series of observations on free
cell-formation without visible nucleus in Alga3 and aquatic
Fungi, especially in Bryopsis, Codium, Anadyomene,
Acetabularia, Dasycladus, Conferva glomerata (lacustris
et marina), and AcJdya prolifera. I regard it as an ab-
normal cell-formation, because it mostly occurs in older
cells, and frequently simultaneously with the death of
the rest of the cell-contents, or precedes this. If this
mode of cell-formation occurs in healthy cells, containing,
in addition to colourless^ transparent contents, and homo-
geneous and granular mucilage, which is especially mani-
fest as the mucilaginous layer investing the internal surface
of the wall, also chlorophyll-globules and starch-granules,
which lie principally in the mucilaginous layer, at first
we see minute, homogeneous, colourless mucilage-globules,
the diameter of which is about '002-'004 of a line. They
become larger, finely granular, acquire a greenish colour,
and a membrane gradually becomes visible upon them.
Subsequently they acquire very various sizes ('OlO-'OGO'"),
and contents which vary much in quantity, since the
chlorophyll sometimes lies upon the wall in smallish
quantity, and sometimes gives the cell a deep green aspect
by its great abundance.

In the above cases the cells originate and are perfected,
without immediately effecting any essential change in the
contents of the parent-cell. But if the parent-cells have

98 VEGETABLE CELLS.

begun to decay, the contents (the mucilaginous layer with
the chlorophyll- and starch-globules) separate from the
wall and lie effused in the cavity of the cell. In this
case the new cells do not usually originate as small,
colourless, homogeneous globules of mucilage, but as
larger globules of the confluent cell-contents, composed
of mucilage, chlorophyll, and amylum. On the very
outside a layer of homogeneous mucilage is always de-
posited, and forms a sharply- defined surface. A distinct
membrane is quickly produced on this surface. The cell-
contents are so arranged within this that the chlorophyll-
and amylum-globules lie at the periphery, while the
middle part of the cavity is filled with transparent fluid.
These cells, whether they be formed within healthy or
decaying cells, originate through abnormal cell-formation,
and stand, apparently, in no necessary relation to propa-
gation ; therefore ordinarily they perish without further
development. In isolated cases, however, they behave
like germ-cells (spores), and are developed into new
plants.

In Plate II, figs. 1 and 2 represent an old utricle of
Bryopsis Balbisiana, wherein abnormal cell -formation is
occurring. In fig 1, a and b, the cell-contents are dead
and beginning to dissolve. They consist of scattered chlo-
rophyll-globules and mucilage-granules. The parts
c and d, on the contrary, possess active living contents,
surrounded by a mucilaginous layer. They are connected
together by a mucilaginous cord : this cord is the rem-
nant of the mucilaginous layer which formerly coated the
part a. In an earlier stage it was thicker, and is now
gradually becoming thinner. Pig. 2 represents the same
portion of an utricle some time later; the connecting
cord of mucilage has disappeared, which has resulted
from its becoming gradually thinner, until it was torn
across, and united perfectly with the two living portions
of the cell-contents, c and d. The former, <?, is now a
defined, ellipsoid portion of contents, altogether free, and
coated over its whole surface with a thin layer of homo-

CELL-FORMATION. 99

geneous, colourless mucilage. The next and most important
change which will occur in it is the appearance of an en-
veloping layer of membrane.

In fig. 3 is also represented a portion of an old utricle
of Bryopsis Balbisiana. With the decaying and dissolv-
ing contents (chlorophyll, starch, and mucilage) occur
small and large cells in various stages. Some are very
small, and do not differ in appearance from a drop of
mucilage (a, a] ; others are somewhat larger, finely
granular, greenish, and already exhibit a delicate mem-
brane (b, b) ; others again are tolerably large and green,
with a distinct membrane (c, c).

Under the head of free cell-formation without a visible
nucleus, the origin of the germ-cells (spores) of the
Zygnemacese (Spiroyyra^ Zyynema, &c.) must also be
mentioned. The facts of the case are well known ; that
two cells unite together (conjugate) ; that the septa of
the point of junction become dissolved ; that the contents
of the two cells separate from their walls, and, either in
the cavity of one of the two cells, or in the tube connect-
ing them, become agglomerated into a globular or ellip-
soidal mass, which becomes the germ-cell. All that
observation shows, in reference to the last proceeding, is,
at first, a mass of green cell-contents, with a definite out-
line, and subsequently the same mass of green contents
inclosed in a membrane. There may be two hypotheses
as to the formation of this membrane : 1, that it origi-
nates in the place where it first becomes visible on the
surface of the contents ; 2, that it originates as a minute
cell in the interior, and, as it increases in size, gradually
absorbs the contents.

In favour of the first assumption speaks, in the first
place, the analogy with the abnormal cell-formation in
Bryopsis, Conferva, &c., where the membrane is in like
manner formed over the surface of the contents. More-
over, an additional support is derived from the circum-
stance that we can see nothing of any little cell in the
interior, or of the alterations in the contents (solution and

100 VEGETABLE CELLS.

reorganization of the solid structures, such as chlorophyll
and starch), which must necessarily be connected with it.
But that hypothesis is placed beyond all doubt by the
following fact. It is well known that the union of two
cells, and the mixture of their contents, do not always
take place. Sometimes a cell produces a germ-cell from
its own contents, when it either forms no conjugative
branch, or when this does not meet with another with
which it can become united. In the latter case the con-
tents separate from the interior of the wall, and move
toward the blind prolongation. Arrived there, any fur-
ther advance being prevented, they become transformed
into a cell, and indeed exactly of the form which they
possess at this time : this happens in Zygnema stdlinum
(pi. II, fig. 4). Sometimes it happens that two cells com-
municate by a connecting tube, but the contents of the
two cells do not become united. Then two germ-cells
originate, one of which is ellipsoidal or globular, while
the other exhibits the form possessed by the contents,
which had already begun to move before they came to a
state of rest and cell-formation (pi. II, fig. 6, in Zygnema
stellinum). In both the cases here described, it must
inevitably be assumed that the membrane originates on
the surface of the contents. If these cells were formed
of minute size in the interior, they must, in their ulterior
development, retain their original globular or ellipsoidal
shape, like all cells which originate and become developed
in a free condition.

Schleiden* is inclined to attribute a different mode of
origin to the germ-cells of Spirogyra, as he says : "In
the already irregularly agglomerated cell-contents, I
almost always found a delicate cell, which I cannot but
regard as the true spore, around which the green and
granular mass is merely applied, forming a false mem-
brane around it, or which gradually absorbs this mass
into its interior. Perhaps the Cytoblast

* Gmndziige der wiss. Botanik, ii, p. 31 (first edition).

CELL-FORMATION. 101

is the producer of the proper spore-cell. " I suspect that
Schleiden has observed a peculiar and enigmatical cell-
or utricle-formation, which occurs in Spirogyra, and of
which I shall hereafter take especial notice. This cell is
neither the rudiment of a germ-cell, nor the germ -cell
itself, nor is it produced from the nucleus of the cell. It
appears to me that the cell-nuclei which lie in the centre
of the cells of Spirogyra do not play any important part
in the origin of the cell. They are often visible until the
cell-contents accumulate into a ball, then they disappear ;
but they are by no means hidden in the contents, since
if they are still visible in the later stages, they are on the
surface of the green contents. I have therefore no doubt
that the nucleus of the parent-cell becomes dissolved
before the contents are transformed into a germ-cell.
Besides, it is not evident what two nuclei should do in
the formation of one cell, each conjugating cell having
one nucleus. In pi. II, fig. 5, some cells of Spirogyra
quinina are figured, in which germ-cells are formed with-
out conjugation. In two of them the nucleus is seen on
the surface of the contents ; in the third it has dis-
appeared.

I mentioned also another free cell-formation, without
visible nucleus, where the observation in like manner
affords a tolerably certain conclusion. This is the origin
of the sporangia in AcUya prolifera. I have already
alluded to this point in the first part of this essay,* in
reference to the so-called division of cells. Aclilya furnishes
remarkable facts, both on parietal and free cell-formation,
so that I will give some figures of it. In the clavately
swollen ends of a branch are formed usually one, more
rarely two or three sporangia. If there be two or three
(PL III, figs. 5 and 6), these originate by free cell-for-
mation ; if there be merely one, it originates sometimes
through free (fig. 3), sometimes through parietal, cell-for-
mation (fig. 7). When the sporangia originate by free

* Ray Society's Translation, p. 279.

102 VEGETABLE CELLS.

cell-formation, the first process is the formation of one or
more heaps of mucilage-granules, according as one or
more cells are to be produced. At first these heaps have
no definite boundary, but pass insensibly into the remain-
ing mucilaginous contents of the cell. Subsequently the
outline becomes more sharply defined, they are surrounded
by ray-like circulation-filaments (fig. 1). The outline
becomes still sharper, and then but a few filaments
remain (fig. 2). At length the surface of the heaps of
granules becomes smooth, and now an inclosing mem-
brane gradually makes its appearance (fig. 3). Has this
membrane originated on the periphery of the granular
accumulation, or has it been formed in its interior as a
minute cell, and in growth gradually absorbed its contents
from without? I am compelled to regard the former
view as correct, because nothing can be seen of the latter
process, even in very transparent heaps of granules. That
the membrane originates on the surface of the subse-
quently inclosed contents is in the highest degree pro-
bable, from the analogy with the parietal cell-formation in
the same plant ; for while a free sporangium is forming in
the interior of one clavate branch, in others the whole
clavate extremity becomes a sporangium (fig. 7). Here
the cell is not formed in the interior as a little free cell,
but its walls originate in the very place where they first
become visible. Now the two facts support each other,
and in such a way that, taken together, they prove the
origin of the membrane on the surface of the contents.
That in the apparent formation of a mere septum, a perfect
cell actually originates,, lying in contact with the wall of
the branch (not a mere wall), is shown by the analogy with
the second case, where a perfect sporangia! cell is formed
at some distance from the wall of the branch. That in
free cell-formation the membrane originates on the surface
of the contents (not in the interior of them, around a
nucleus, or in any other way), is shown by the analogy
with the first case, where the cell-membrane is formed in
like manner on the periphery of the contents. The dis-

CELL-FORMATION. 103

tinction lies solely in this : that in the first case the whole
contents of the extremity of the branch, in the second
only a part of them, are converted into a cell.*

b. With a parietal nucleus.

Free cell-formation with a visible nucleus I have
hitherto only observed with certainty in the embryo- sac
of the Phanerogamia. The cell-formation in the pollen-
tube is not yet clear to me ; of those cells which form the
embryo and the suspensor, I rather imagine that they
originate through parietal cell-formation ; of those cells
which Schleiden explains as transitory cells, I am still
doubtful whether they are actual cells, or not rather mere
nuclei.

When the formation of the endosperm-cells takes place
in the embryo-sac, the fluid contains the following struc-
tures :

1. Cells with granular contents, and a perfect nucleus,
with evident nucleoli.

2. Mucilage- granules.

3. Perfect nuclei, with evident nucleoli.

4. Minute homogeneous globules of mucilage.

5. Larger homogeneous globules of mucilage, with a
globular cavity.

6. Larger homogeneous globules of mucilage, with a
smaller concentric or excentric ring.

* In the sporangium which has originated in the above ways, two kinds
of cells may be formed, either larger, immoveable, globular spores, with a
tough membrane, or smaller oval cellules, with a delicate membrane, which
move about actively either already inside the parent-cell or, and especially,
after they have been set free. To this end the sporangium grows out into
one or more processes (figs. 4, 6, 8), which open at the point and allow the
cellules to escape. I formerly believed that even a third kind of cell origi-
nated in the sporangium, namely, smaller cells with delicate walls like the
moving cellules, but exhibiting no motion, and capable of germination.
Whether this third kind actually exists, or whether they are identical with
the moving cells, and again, whether these are capable of ^ermination, I
must leave open, since the latter has been stated to be the lact by several
observers, till further researches have been made.

104 VEGETABLE CELLS.

7. Mucilage-globules, of equal size with the preceding,
partly homogeneous, partly very slightly granular, with a
cavity and a nucleolus contained in it,

8. Like 7 ; but with several cavities, and as many
nucleoli.

9. Like 7, with one or more solid nucleoli, and no
cavity.

10. Larger homogeneous or slightly granular mucilage
globules, of clearer, more transparent substance, with or
without cavities, with or without nucleoli.

11. Clear vesicles, of the size of 10, with or without
nucleoli.

12. Mucilage-globules, like 10, or utricles, like 11,
surrounded by a thin layer of homogeneous mucilage,
which is usually thinner on one side than on the opposite.

13. Like 12 ; the mucilaginous layer has considerably
increased in size on one side, become somewhat granular,
and displays a very indistinct membrane on its periphery

14. Like 1, namely, distinctly perfect cells, with per-
fect nuclei and distinct nucleoli.

These are the principal and most frequent forms of
organization that are found in the fluid of the embryo-
sac, and which are of importance in the processes which
go on. A quantity of intermediate stages between the
various forms here enumerated are rather calculated to
confuse the vision and judgment than to make a pro-
cess clear through a complete history of the gradual de-
velopment. Observation of these matters has to strive
against two obstacles, which appear to me almost insur-
mountable : one depending on the fact that partial changes
ensue very rapidly in the solid portions of the contents,
especially when water is applied the other, that the dif-
ferent stages of development are all mixed together, and
thus, without the help of extraneous characters, are ne-
cessarily judged of merely by their own individual aspect.

Schleiden, it is well known, has interpreted the pheno-
mena in this way : that a number of mucilage-granules
become confluent to form nucleoli ; that the nucleoli

CELL-FORMATION. 105

become conglomerated with other mucilage-granules, so
as to produce a nucleus, and that around the nucleus is
formed a membrane, which retreats from the nucleus on
one side through the absorption of fluid.

Now as to the mucilage-granules, which lie in the
effused fluid, I believe that they are derived, in great part,
from the destroyed endosperm-cells, and that they do not
at all contribute to the formation of the nuclei. In the
embryo-sac, where cell-formation is commencing, I often
find but little, often no granular contents ; these increase
with the fuller development of the endosperm-cells, and
with the cessation of cell-formation. This circumstance
already makes the origin of the nuclei from mucilage-
granules very improbable.

Again, I was never able to observe a confluence or
conglomeration of mucilage-granules, and the production
of a nucleus therefrom. The solid portion of the cell-
contents certainly concreted into larger or smaller masses,
especially when water was added. Sometimes only a few
mucilage-granules, sometimes mucilage-granules and nu-
clei, sometimes large portions of the firm granular contents
became united ; a homogeneous mucilage was always the
connecting medium. This, however, does not happen
through any organic process, but because the mucilage
coagulates. The conglomerated masses do indeed ex-
hibit, like all coagulating mucilage, a definite outline,
and, when they are small, sometimes present a striking
resemblance to nuclei or young cells. But they are often
actually seen to be formed, in the same way that they
have been formed here, when the mucilage of any cell
flows out into water.

Besides the mucilage-granules, we must likewise leave
out of consideration, in cell- formation, the clear vesicles.,
which might easily be taken for nascent cells, but which
are found in almost all homogeneous and rather fluid
mucilage. They appear to me to be minute drops of
some watery fluid (water?), which have separated from
thicker, mucilaginous fluid of the cell. In most cases the

106 VEGETABLE CELLS.

mucilage exhibits an equal density up to the border of the
drop of water, so that where the water commences the
mucilage appears as if cut off from it. Sometimes it is
denser at the borders of the drop of water, and seems to
form a proper membrane around it. Probably this is the
consequence of a lengthened action of the water upon the
mucilage. In any case the watery vesicles go no further ;
certainly undergo no organic changes.

All perfect nuclei lying in the fluid must also be ex-
cluded in the consideration of cell-formation. They are
wholly absent at first, and multiply in proportion to the
number of fully-developed endosperm-cells contained in
the embryo-sac. They are the nuclei of these cells, and
become free, with the granular contents, through their
destruction. These nuclei are very apt to lead the ob-
server astray, since they are not capable of the subsequent
stages of development.

Small, globular drops of perfectly homogeneous mucilage,
with defined outline, are absolutely constant phenomena in
the process of cell-formation in the embryo-sac. They
vary from '001 -'004 of a line, and are distinguished from
the mucilage-granules, even in the earliest condition,
by a perfectly globular form and smooth surface. The
mucilage-globules are never absent ; they always con-
stitute the first stage of cell-formation (pi. II, fig. 7, a,
b, c; 8, a). They exactly resemble the mucilage-globules
which represent the nascent germ-cells in Valonia and
other Algae, in Lichens and Fungi.

A more advanced stage is represented by the larger
mucilage-globules, in which a smallish ring is inclosed,
ivhile the whole olobule possesses an uniform consistence

/ JL */

(fig. 7, d, e, /). The inclosed ring may appear clearer
(fig. 9, a] or denser (fig. 9, b) than the outer part of the
globule. There can be no doubt, and the further deve-
lopment also confirms it, that the mucilage-globule is a
cell-nucleus, the inclosed ring a nucleolus. How have
the nucleus and nucleolus originated ? This is a question
which I cannot answer from direct observation on cell-

CELL-FORMATION. 107

formation in the embryo-sac. Other grounds, however,
which exist in the phenomena exhibited in the propaga-
tion of the nuclei, render it probable, in my opinion,
that the nucleolus originates first, and the nucleus subse-
quently, around it.* In that case, those little homogeneous
mucilage-globules, first visible, would have to be regarded
as the nucleoli.

At this stage of development we find, as a rule, a
homogeneous nucleus inclosing a homogeneous nu-
cleolus. Through injurious external influence, a hollow
space round the nucleolus is produced. This change oc-
curs during the examination. Fig. 9 represents such a
nucleus, immediately after the fluid of the embryo-sac
had been brought under the microscope. It soon ac-
quired the aspect d (fig. 9). This appearance of a hollow
space round the nucleolus is observed both when the fluid
of the cell is diluted by the addition of water, and when
it is brought on the stage without water, and becomes
denser through evaporation. The cause of this pheno-
menon appears to me to lie in the fact, that through the
action of the slightest possible unfavorable influence from
without, the mucilage of the nucleus, as well as that of
the nucleolus contracts, and they are thus separated from
each other. If the action be at all strong, the nucleus
and nucleolus contract into a dense, solid body. Care
must therefore be taken not to mistake abnormally altered
nuclei, such as are drawn in fig. 9, d, for cells with nu-
cleus and nucleolus, with which they may very readily be
confounded.

Other nuclei contain two, three, or four nucleoli (fig.
7, g \ fig. 8, c\ fig. 9, e). These nucleoli are also some-
times of the same consistence as the nucleus, sometimes
they are denser, at others not so dense. In abnormal
alteration of the nuclei, a hollow space presents itself to
notice around each individual nucleolus. Fig. 9, e and/,
represent such a nucleus before and after alteration.

* See the Essay on the Utricular Structures iu the Contents of the
Vegetable Cell.

108 VEGETABLE CELLS.

These forms with two, three, or four nucleoli seem to prove
that the nucleus generally originates before the nucleoli.
Since no distinction is evident in the nucleoli, one is in-
clined to regard them all of equal value ; and since some
of these nucleoli certainly originated after the nucleus,
one is led to assume the same of all. I believe, how-
ever, the assumption that one of these nucleoli originated
before the nucleus and produced it, and that the rest were
formed subsequently, may be warranted by the analogy
with cells. In cells also we sometimes find several nuclei,
a primary and one or two secondary nuclei.* Therefore,
since both the life of the nucleus with the life of the cell,
and the relation of the nucleolus to the nucleus with that
of the nucleus to the cell, exhibit so great analogy, we
may reasonably assume, that when a nucleus contains
several nucleoli, one of them is the primary, which ori-
ginated prior to the nucleus, while the rest are secondary
nucleoli formed within the nucleus.

The nuclei have now attained a definite magnitude.
Their membrane may sometimes be seen indistinctly on
their periphery. Their mucilaginous contents are either
homogeneous or very slightly granular, of variable density.
The substance of the nucleoli is generally denser than that
of the nuclei, sometimes of equal, at others of a less, de-
gree of density, so that in the last case the nucleoli appear
as clearer spaces in the denser contents of the nuclear
vesicle.

Subsequent stages exhibit nuclei, like those just de-
scribed, with a thin layer of homogeneous mucilage. This
layer is usually much thinner on one side than on the
opposite (fig. 8, d). More rarely it is equally thick on
all sides ; which may be ascertained from the fact, that, in
rolling, the nucleus always retains its central position (fig.
8, e). The mucilage is perfectly homogeneous, distinctly
defined externally, and bounded by a somewhat darker
line. As yet I can discover no membrane. The mucilage

* See Part I of this Essay. Ray Society's Translation, pp. 242 et seq.,
and pp. 247 et seq.

CELL-FORMATION. 109

is mostly somewhat more dense than the substance of the
nucleus ; sometimes it is brighter than the latter.

The mucilage surrounding the nucleus now continually
increases in quantity; the nucleus is always distinctly
visible, situated at the periphery (fig. 8,/). There are,
however, isolated exceptional cases here, where the nucleus
is free, and more or less removed from the circumference.

The mucilage then becomes evidently granular, and a
distinct membrane is to be made out upon its surface
(fig. 8,y). The latter frequently becomes visible even when
the contents are still homogeneous. In this stage, as in
the preceding, the nucleus is mostly less dense than the
surrounding mucilage, and therefore appears like a clearer
space. The cell is now visibly formed. Whether the
membrane first originates at the time it becomes visible,
or had originated at a still earlier period, can scarcely be
ascertained by observation. The latter appears to me
probable.

According to Schleiden, the cell-membrane originates
immediately on the surface of the nucleus ; it absorbs
watery fluid by endosmose, and expands ; the nucleus re-
mains attached on one side of the cell ; the cell is a fine,
transparent vesicle ; its contents a watery fluid, and ap-
pear merely as a hollow space between the nucleus and
the granular mucilage of the embryo-sac, which is pushed
backward by its expansion.* Schleiden adds, that the
cells become wholly dissolved in a few minutes in distilled
water, so that only the nuclei remain. I confess that I
have never seen such very clear and transparent cells.
In most cases I find the contents of the cells more dense
and darker than those of the nucleus ; rarely clearer, but
even then always either homogeneously mucilaginous or
finely granular. In like manner, I did indeed see various
effects on the young cells and their contents produced by
the action of water, but never a solution.

* Miiller's Archiv, 1838. [Schleiden has changed this opinion in the
latest edition of his ' Principles of Botany' 1849). See Appendix to Dr.
Lankester's Translation. A. H.]

110 VEGETABLE CELLS.

I will here mention another phenomenon which I ob-
served a few times. I do not think that Schleiden has
allowed himself to be deceived by this ; yet others, who
may also observe it, may easily suppose it to be the process
of cell-formation as described by Schleiden, to which it
bears great resemblance. Nuclei which have attained a
considerable size, absorb water by endosmose. The
membrane of the nucleus detaches itself on one side from
the contents, the interspace becoming filled with water.
The comparison with a watch-glass is here not inapt. In
some cases, the membrane continues to expand, suddenly
bursts, and disappears. In others, this expansion goes
on to a variable extent, and ceases when the force of the
endosmose and the elasticity of the membrane come into
equilibrium. The membrane remains visible and is not
dissolved. (See fig. 9, i, k, I, m ; in h, i, k the same nu-
cleus is represented previous to the action of the water,
and in two stages after the operation of endosmose has
begun.)

If the nuclei contain only one nucleolus, one may
readily suppose them to be cells, and the hollow space
round the nucleolus to be the nuclear vesicle (fig. 9, i, fc).
Those forms, with two or three nucleoli (fig. 9, /), how-
ever, prove that it is inside the nuclei that the hollow
spaces have been formed through the action of water, as
is the case in the nucleus represented in fig. 9, c-g. The
contents of the nucleus are pretty sharply defined where
they join the water. The outer circumference of the
nucleus exhibits a dark line, which is formed by the de-
licate membrane lying close upon the contents, and which,
therefore, is lost when this membrane has become detached
from the contents (fig. 9, h, /, 0). These facts furnish a
new proof that the nuclei possess a membrane and are
vesicles.

The young cell appears at first as a layer of mucilage
surrounding the nucleus. Subsequently a membrane
becomes visible on the surface of the mucilage. Conse-
quently the cell is at first, besides the nucleus, quite filled

CELL-FORMATION. Ill

with mucilage. When the cell expands more, it becomes
hollow in the interior. The mucilage remains on the wall
as a thin layer, and forms a coating over the whole inter-
nal surface. This is the mucilaginous layer which I have
described in the Alg8e,*and which Mohlf has named the
" primordial utricle" regarding it as a structure proper
to all cells. The mucilaginous layer is usually thicker at
the place where the nucleus of the cell lies, than over the
rest of the wall. Not unfrequently the nucleus lies
wholly imbedded in the mucilaginous layer. Schleiden
thought that it was inclosed in a duplicature of the cell-
wall. But this is certainly incorrect, and is best refuted
by the fact, that the nucleus may become detached, with
the mucilaginous layer, from the membrane of the cell.

Mohl conjectures that the first thing formed around a
nucleus is the primordial utricle, and that the membrane
does not originate until after this. The statements in
regard to this are, however, too vague, and made without
consideration of cell-formation in the embryo-sac, so that
no minute discussion is necessary for the criticism of the
phenomena in question. I merely remark that I see the
origin of the mucilaginous layer (primordial utricle) on
the internal surface of the cell in the endosperm-cells, in
the same manner as in all other cells where new contents
are formed. The mucilage at first fills the whole cavity
of the cell, and subsequently merely lines the walls as a
thin layer. The mucilaginous layer is, therefore, a
secondary phenomena in the origin of the cell-contents.

c. Of free cell-formation as a general law.

Now that I have separately brought forward the facts
which are at my command respecting free cell- formation,
I will briefly collect and compare them, in order to

* See Part I of this Essay. Ray Translation, p. 268.
t Botan. Zeit. 1844. (Translation in Taylor's Scientific Memoirs, vol. iv,
p. 91.)

112 VEGETABLE CELLS.

deduce a general law, and, in addition, to point out
some essential differences within the bounds of this law.

The phenomena which accompany free cell-formation
may be seen with the greatest certainty in the origin of
the germ-cells of Zygnema, the sporangial cells of Achlya,
and the larger cells produced by abnormal formation in
Bryopsis, Conferva, and other Algse. Here a portion
(in Zygnema the whole) of the contents becomes isolated,
acquires a globular or ellipsoidal form, and produces a
perfect closed membrane all over its external surface.
In the prevalent opinion on free cell-formation, it has
been hitherto assumed that the membrane originates
around a nucleus. In the cases mentioned here there
cannot be any doubt that the membrane originates around
the contents. It was further assumed that the membrane
was formed first and the contents subsequently. Here,
however, the contents are decidedly primary and the
membrane secondary.

The mode of origin of the endosperm-cells in the
embryo-sac comes next to these facts. Observation
shows here, first a nucleus, then a layer of mucilage which
surrounds this nucleus, lastly, a distinct membrane which
incloses the mucilage together with the nucleus. The
phenomena accessible to observation leave the place and
epoch at which the membrane originates undecided. The
assumption that it is formed immediately around the
nucleus is, therefore, still within the bounds of possi-
bility. But it would appear to me uncalled for and
superfluous, since no fact nor analogy are in its favour.*
If we assume, on the other hand, that a definite quantity
of mucilage collects around the nucleus, and that the cell-
membrane originates around this mucilage, the assump-
tion on the one hand fully meets the appearances in the

* Schleiden's observations certainly are contradictory to my representa-
tion and interpretation, and these points are still open. Formerly I thought
that I also saw cell-formation round the nucleus. I have studied the pro-
cesses occurring in the embryo-sac repeatedly, for several years, and in
different plants ; with more accurate observation, however, I can no longer
find any condition which perfectly agrees with Schleiden's representations.

CELL-FORMATION. 113

case in question; and on the other, connects itself,
through analogy, with other facts, as in the cell-formation
already mentioned, in Acldya, Zygnema, and other Algge,
as well as with the parietal cell-formation, where the
membrane is likewise produced on the surface of the
contents.

The third type of free cell-formation still remains, in
the origin of a number of germ-cells of Algae, Fungi, and
Lichens. Here observation at first discloses to us ex-
tremely small globular masses of mucilage. They become
larger, granular, and at last an inclosing membrane may
be made out. Actual experience is again insufficient to
furnish a certain determination of the question when and
how the membrane originates. It seems to me safest
to take the simple explanation of the facts which is first
suggested by what we see, especially since they can in
this manner be most readily brought into agreement with
the other facts. I therefore assume here, that the mem-
brane is formed round a collection of the mucilaginous
contents. Whether this happens earlier or later, seems
to be all one ; but it is probable that the membrane
actually exists some time before it is distinctly visible.
The observations on free cell-formation therefore require,
they partly allow, the assumption, that the membrane is
produced on the surface of the contents. This hypothesis
must, since no facts and no analogy are opposed to it, hold
as a universal law. The contents, therefore, are primary,
in the cell, the membrane is only secondary.

In the production of the endosperm-cells a nucleus
exists. The nucleus originates first. Subsequently, a
layer of mucilage accumulates on its surface. Probably
it is the attractive force of the nucleus which draws to it
a portion of the contents of the parent-cell. That the
nucleus does possess such a power is known from many
facts. In almost all cells in which a nucleus exists, a
portion of the contents of the cell become collected on its
surface. Consequently, we may define cell-formation in
the embryo-sac in the following way : A nucleus origi-

114

VEGETABLE CELLS.

nates, which attracts a small portion of the mucilaginous
contents, and becomes wholly clothed by it : on the surface
of the mucilaginous layer the cell-membrane originates.

In the other cases of free cell-formation, nothing can
be seen of nuclei during the whole process. Either no
nucleus really exists, or it withdraws itself from vision,
through relatively minute size, or agreement of density
with the density of the contents of the young celL In
the first case we must assume, that a definite portion of
the contents of the parent-cell may independently become
individualized and converted into a cell. In the second
case, the process would be the same as in the endosperm-
cells. In my opinion a rigid distinction must be made
between normal and abnormal cell-formation, in solving
this question. By normal cell-formation I understand
such as is necessarily connected with the vegetative and
reproductive processes in a plant, and which always pro-
ceeds according to laws definitely laid down for each
plant. By abnormal cell-formation, on the other hand,
I understand such as is not directly and necessarily con-
nected with the vegetative and reproductive processes,
and must always be regarded more or less as produced
by the action of external influences interrupting the
regular course of the life of the cell.

As to the abnormal cell-formation in the cells of Algae,
I am convinced that no nuclei ever take part in it. For
this kind of cell-formation is united by every possible
intermediate stage, with a process in which the formation
of nuclei cannot enter into the question. In the first
part of this Essay* I have mentioned a partial formation
of membrane where, in consequence of disturbing external
influences, the mucilaginous layer is retracted in places
from the cell-wall, and becomes coated with new pieces
of membrane. This process is of course independent of
the influence of any nucleus. I, at the same time, stated
that the mucilaginous layer sometimes becomes detached

* Ray Translation, pp. 268 et seq.

CELL-FORMATION. 115

over large surfaces, or even entirely (as in Bangia) from
the cell- wall, and produces a membrane on its surface ;
moreover, that sometimes the mucilaginous layer divides
into separate portions, and produces several perfect cells.
It is clear that in this case, again, we cannot think of the
formation of a nucleus for the production of the cells.
These facts stand in direct connexion with those above
described in abnormal free cell-formation. The mucila-
ginous layer separates from the wall, divides, and forms
several larger or smaller cells. These vary in size and
contents, from, large cells containing chlorophyll and
starch, to the most minute cells inclosing merely homo-
geneous mucilage. This mode of cell-formation varies,
besides, from that condition in which the whole cell-con-
tents partly form new cells, the remainder being dissolved,
to that in which merely a small portion of the contents
forms one or more small cells, while the remainder is
unaltered. These transitions show that all the pheno-
mena of abnormal cell-formation are related to one law,
and that since the formation of nuclei is inadmissible in
some cases, it must not be assumed in the rest.

In those cases where free cell-formation takes place
normally, as in the origin of the germ-cells in Algae,
Lichens, and Fungi, and of the sporangium in Achlya,
we can see nothing of a nucleus. The fact, as presented
directly to us, with the assistance of the amplifications
our present optical instruments effect, may in the same
way be most simply explained thus : that larger or smaller
portions of the contents at once become individualized,
and acquire a membranous investment. But the circum-
stance that a nucleus may sometimes be perceived subse-
quently, in the germ-cells which have originated by free
cell- formation, seems to me to speak against this. I have
seen it in Erysibe, AcJdya, Peziza, Coleoclicete, &c.; and it
appears to me to be also present in the germination of
the Zygnemacese. Two explanations may be given as to
this nucleus. Either it is a primary nucleus, around
which the cell originated, or it is a secondary nucleus

116

VEGETABLE CELLS.

which has been formed as an after-growth in the cell.
We are acquainted with such secondary nuclei in the
parent-cells of spores and pollen-granules, and in the
spore- and pollen-cells themselves.* In these, however,
there is also a primary nucleus. The two nuclei lie side
by side in their cell ; or the primary nucleus becomes
dissolved when the secondary is produced. In the cases
referred to, the primary nucleus is parietal, the secondary
free. Now, as regards the nuclei in the germ-cells of the
Algae, Lichens, and Fungi, in some I found a distinctly
parietal position. I conjecture, therefore, already on these
grounds, that they are primary nuclei, and, consequently,
that a nucleus exists in the origin of the germ-cells. This
conjecture is supported by another reason. In the section
on the nucleus,! I demonstrated that if any conclusion
from analogy at all be permitted, it must be assumed that
every vegetable-cell possesses a nucleus, at least in the
early stages of its existence. It is, therefore, in the
highest degree probable that a nucleus is present in the
cell originating by normal cell-formation, and this at the
actual time of its origin. This argument is still further
borne out by the fact that sometimes, in plants where no
nucleus is visible in the formation of the germ-cells, all
the succeeding cells are developed under the influence of
nuclei. Now it seems to me very improbable here that
the vegetative cell-formation should take place through
nuclei, and the reproductive cell-formation without, that,
consequently, the lower cell-formation should present
greater complexity, and the higher be the more simple.
It is further to be remarked, that the assumption that
germ-cells originate like endosperm-cells around nuclei,
may be connected with the visible phenomena without
stretching any point. In both places, globules of muci-
lage first present themselves, which gradually enlarge, and
at last appear as granular cells. In the endosperm-cells,
the nuclei may be distinguished and recognised as such,

* Part I, Ray Translation, p. 247. $ Ibid., pp. 219 et seq., and 246.

CELL-FORMATION . 117

at a very early period. In the germ-cells, the nuclei are
distinct, as such, sometimes never, sometimes only in the
fully-developed condition of the cell. Since the nuclei
can be distinguished sufficiently early in the endosperm-
cells, it is also possible to observe their relation to cell-
formation. Since the nuclei of the germ-cells cannot be
distinguished until a late period, their relation to cell-
formation cannot be made out by direct observation. This
is the more conceivable, that the endosperm -cells are many
times larger than the germ-cells, and their nuclei than
nuclei of the latter. Besides this, endosperm-cells do also
occur in which the presence of a nucleus, and its coopera-
tion in the cell-formation, cannot be distinctly seen. This
is especially the case when the contents of the nucleus
and those of the young cell are homogeneous and of equal
density. What I have stated is, I think, sufficient reason
why the apparent want of nuclei in the free formation of
germ-cells cannot be maintained as a proof of an actual
absence of these bodies.

Normal free cell-formation thus comprehends, according
to my views, the following essential and regular periods :
A nucleus originates in the contents of the parent-cell.
This accumulates on its surface, by attraction, a greater
or smaller quantity of the contents of the parent-cell,
which, at least at the periphery, consist of homogeneous
mucilage. This portion of the contents becomes coated by
a membrane over its entire surface.

The variations which occur within the limits of this
identity, refer either to the nature of the portions of con-
tents which become individualized, or the relation of the
secondary cells to the parent-cell. The portion of con-
tents which become isolated, exhibit chemical and mor-
phological variations. They are composed either of
homogeneous, colourless mucilage ; of homogeneous
colourless mucilage mingled with colouring matters; of
granular mucilage, or granular mucilage mixed with
colouring granules and starch-globules.

The relation of the secondary cells to the parent-cell

118 VEGETABLE CELLS.

depends either on the share which the parent-cell bears in
the production of the secondary cells, or on the physiolo-
gical similarity or dissimilarity which exist between the
parent-cell and the secondary cells. In reference to the
first point, it is of importance whether the whole contents
of the parent-cell, or only larger or smaller portions of the
contents, are converted into secondary cells, and in what
number these are produced in the parent-cell. In refer-
ence to the second point, either perfectly similar or dis-
similar secondary cells may be formed in the parent-cell.
Physiologically similar secondary cells are produced in
Hcematococcus and Chlorococcus ; all other cells originating
through free cell-formation, are more or less different in
physiological respects from the parent-cell.

The question still remains, to what extent is free cell-
formation met with in the vegetable kingdom ? In the
first part of this essay,* I limited parietal cell-formation
to several families of Algae, and the special parent-cells
of the four-spored Cryptogamia and the Phanerogamia.
For all the examples there enumerated, I had observations
which more or less bore out the statement. Free cell-
formation, on the other hand, I was acquainted with in
the endosperm-cells and in the germ-cells of Algae, Fungi,
and Lichens. For all other cases, especially for all vege-
tative cells of four-spored Cryptogamia and the Phane-
rogamia, the decision depended partly on analogy, partly
on some observations of Schleiden's, and of my own. I
therefore concluded : In parietal cell-formation the nu-
cleus is, as a rule, central ; in the free cell-formation in
the embryo sac, the nucleus is lateral. It is probable from
this, since the cells of the higher Cryptogamia and the
Phanerogamia possess a lateral nucleus, that they origi-
nate by free cell-formation. The secondary cells were,
moreover, figured actually free within the parent-cell, in
certain cases, by Schleiden and by myself. I was satisfied,
therefore, that the cells with lateral nuclei originated free.

Observation and reflection have since led me to a

* Ray Trans., p. 292.

CELL-FORM ATION . 119

different conclusion. In the very first place comes the
question, what import may be attributed to those obser-
vations which represent the secondary cells free within
the parent-cells, as Schleiden's in the germinating spore
of MarcJiantia* in the hairs of the ovary of Lupinus,^
in the terminal shoot of Opuntia,\ in the pollen-tube, and
in other cells, and my own in the apex of the root of
Lilium. || As to my own earlier investigations, they be-
long to an epoch (autumn 1841, and spring 1842,)
when I was yet unacquainted with the mucilaginous
layer. When I discovered it, in the summer of 1842,
at Naples, in Algae and Florideae, and subsequently re-
found it in other vegetable-cells, I soon became convinced
that my earlier researches were doubtful, since that which
I had held to be the cell-membrane might in every case
have been the mucilaginous layer.

With regard to the facts published by Schleiden, I
likewise believe that the explanation given by him is by
no means placed beyond all doubt. In the first place,
cell-formation in the pollen-tube cannot prove anything
for other cell- formation ; since here two thoroughly dif-
ferent kinds of cell-formation, one of which is transitory
while the other produces the embryo, have not yet been
nearly sufficiently discriminated. The cell- formation in
the spore of Marchantia, I must, from certain researches
of my own on the germination of the Hepaticse, hold to
be incorrect ; and conjecture that a confusion has occurred
with the accidental formation of utricles, such as some-
times happens in cells. With respect to the rest of the
observations published by Schleiden, it must be borne in
mind that he also makes no distinction between cell-
membrane and mucilaginous layer ; so that the interpre-
tation is always possible, that the apparently free secondary

* Miiller's Archiv, 1838, Tab. iii, figs. 19, 20.
- Acta Ac. C. L. C. Nat. Cur., vol. xix, p. 1, pi. x, fig. 38 e.
% Mem. de 1'Acad. Imp. d, Sc. de St. Petersbourg, VI Serie, vol. iv,
pi. viii, fig. 9.

Grundz. 2d Ed. pi. i, figs. 12 and 16.

Linnaja,

2d Ed. pi. i, figs. 12 and
1842, PI. ix, figs. 30, 31.

120 VEGETABLE CELLS.

cells have been merely the contents of secondary cells
contracted through external influences. Two reasons
especially strengthen this opinion in my mind. In one
case Schleiden figures three secondary cells in one parent-
cell. Now, as I shall hereafter show, it is an universal
law that in the formation of the cells of tissues only two
secondary cells are formed in each parent-cell. If, then,
one wall have been overlooked in consequence of the
alteration of the contents, so readily may the other.
Schleiden further says that he has often seen two cells
inside one cell,* especially after the application of nitric
acid. This is in contradiction with the observation he
made, that the young cell-membranes are wholly dissolved
in distilled water, in a very short time.f If this will
happen in water, it will certainly happen much more
quickly in nitric acid. It is in the highest degree pro-
bable from this, that Schleiden saw merely the contracted
contents, not the young cell and its membrane; and,
moreover, in that condition in which the action of the
acid has already made the delicate membrane of the
secondary cells indistinct, while the older and firmer
membrane of the parent-cell still remains visible.

Since then a better appreciation of the membrane, and
the recognition of the mucilaginous layer, rendered all
earlier observations uncertain, I instituted a new series
of investigations. If the tissue of the higher plants were
really produced by free cell-formation, this could only be
proved by seeing the young free cells, free in uninjured
and unaltered parent-cells. I must confess that I never
have arrived at this. In most cases this is, indeed, no
proof at all of the contrary ; since admitting a free cell-
formation, it would, in my opinion, be unlikely, in the
majority of cases, that the delicate membranes should be
seen in the but slightly transparent mucilage, previously
to their union to form a wall. On the other hand, I be-
lieve that in certain other instances it might possibly have

* Grundz. 2d Ed. p. 202. f Miiller's Archiv., 1838, p. 145.

CELL-FORMATION. 121

happened, that the young free cells should have been seen,
in some when the parent-cells were larger, and filled
with granular contents (as in many hairs when their de-
velopment is beginning, e. g. of Tradescantia) ; in some
when the parent-cells, of considerable size, contained a
homogeneous and tolerably transparent fluid (as in the
young rind and bark of many plants) ; in others when
the parent-cells were tolerably large, and more or less
filled with coloured, homogeneous or granular contents
(as in Caltithamniacea). But in no case could I see the
young free cells inside the parent-cells. The formation
of the cells of tissues in the higher Cryptogamia and the
Phanerogamia, displayed to me nothing but either merely
a dividing wall, or two nuclei and then a wall between
them, or a larger nucleus which divided into two nuclei,
and then a septum formed between these.

In consequence of these researches I have come to the
conclusion, that all vegetative cell-formation is parietal.
This conclusion is supported on the one hand by the facts
observed, on the other by analogy. Among the facts,
some do not contribute to the proof; others make the
parietal cell-formation probable ; a few can be explained
by it alone. In Griffitksia corallina* (pi. Ill, fig. 9), the
parent-cells are very large, as much as * 080 of a line long
and more, and sometimes of almost half that breadth.
The granular contents lie upon the wall ; the interior is
filled with clear, colourless fluid. The cell divides into
two unequal cells, at the apex, by a cross wall (fig. 10) :
an upper cell, small, disc-shaped, and wholly filled
with granular contents (a), and a large lower cell (3), re-
sembling the parent-cell in all its parts. If the two
secondary cells were formed free, the lower, especially,
must be seen during its development, and changes in the
parietal firm contents be perceived, when this became
dissolved in the parent-cell and reorganized in the secon-

* Griffithsia belongs to the Floridete ; the Florideee, however, have not a
natural relationship to the Algse, but with the Hepaticae and Mosses. They
also, like these, possess parietal nuclei in the cells.

122 VEGETABLE CELLS.

dary cell. The appearances are the same here as in the
parietal cell-formation of the Algse. From this indubitable
example we have numberless intermediate stages to those
cases in which no evidence is afforded, either for or
against. The succession of states and the transitions
show, however, that the same explanation must come into
application for all cases.

The conclusion that all vegetative cell-formation is
parietal, is also supported by analogy. The parietal cell-
formation of Algae and special parent-cells occurs, as a
rule, with central nuclei. But to this rule there are some
exceptions, when the parietally originating cells possess a
lateral nucleus.* In the vegetative cell-formation of the
higher Cryptogamia and the Phanerogamia, the nucleus
is, as a rule, lateral ; yet it appears to me that, in ex-
ceptional cases, it may be free also. The appearances in
the Algse and special parent-cells on the one hand, and in
the cells of other plants on the other, are essentially the
same, only they present themselves much more dis-
tinctly in the first, and much more definitely require to
be explained as parietal cell-formation.

Parietal and free cell-formation would, according to my
views, extend through the vegetable kingdom within the
following bounds :

Parietal : the vegetative cell-formation of all plants,
further, the reproductive cell-formation of many Alga and
Fangi. Free : the reproductive cell-formation of most
(not all) plants, namely, the germ-celt, formation of many
Fungi, many Algce, and of the Lichens ; the formation of
the spores inside the special parent-cells in the four-spored
Cryptogamia (?), the formation of the pollen-cells inside the
special parent-cells in the Phanerogamia (/*), and the forma-
tion of the endosperm-cells in the Phanerogamia.

The cells which have originated by parietal cell-forma-
tion possess central nuclei (in most Algae, and generally
the special parent- cells), or lateral nuclei (in most Fungi,

* Part I, Hay Trans., p. 552 ct scq.

CELL-FORMATION. 123

in the Cryptogamia from the Floridece upward, and in the
Phanerogamia) . The cells which have originated by free
cell-formation contain, as a rule, a lateral nucleus, rarely a
central nucleus (the germ-cells of certain Algse).

V. ON CELL-EORMATION IN GENERAL.

Now that I have discussed, in the third section of this
essay, parietal, and, in the fourth, free cell-formation, I
will compare these two processes, and thence deduce a
general law.

In parietal cell-formation the contents of the parent-cell
divide into two or more portions. Around each of these
portions of cell-contents a perfect membrane is produced,
which, at the moment of its origin, is in contact partly with
the wall of the parent-cell, partly with the corresponding
wall of its fellow secondary cell or cells.

In free cell-formation a greater or smaller portion of the
contents becomes isolated, or even the whole contents of the
cell. On the surface of this is formed a complete mem-
brane, altogether free at its outer surface (in contact neither
with the wall of the parent-cell nor those of its fellow
secondary cells.

Cell-formation includes two stages ; the first is the
isolation or individualization of a portion of the contents of
the parent-cell ; the second consists in the origin of a
membrane around individualized portions of the contents.
Cell-formation commences with the first, and is completed
in the second stage. The complete individualization only
occurs for the express purpose of cell-formation. The
formation of membrane, on the contrary, is a universal
phenomenon, proper to the cell generally (not merely at
the moment of its origin). I will first speak of this latter.

In the first half of this memoir, speaking of the cells of
the Algae, I have noticed the mucilaginous layer which
lies on the outermost boundary of the contents, and there-
fore close upon the surface of the cell-wall. According

124 VEGETABLE CELLS.

to Mohl, this is of very general occurrence. From my
own researches, I believe that I may venture to affirm,
that, with the exception of those young cells, in which the
cavity is filled with homogeneous mucilage, the muci-
laginous layer is always present in cells so long as they
retain their vitality. It is well known that the thickening
of the cell- walls is effected by a deposition of new layers.
These new layers are produced between the cell-wall and
the mucilaginous layer. This fact admits of no other
explanation than this : that the mucilaginous layer (or the
contents through this layer) secretes organic, unazotized
molecules which form the new thickening layer.

I have further shown, in the first part of this essay, that
in consequence of disturbing external influences, in the
cells of various Algae, the mucilaginous layer is retracted,
in places or entirely, from the wall, and produces a mem-
brane on the surface, which becomes free, and bounded
externally by water. This membrane is not merely similar
to cell-membrane in outward conditions. It presents it-
self as such in all its characters, so that, in particular, it
has the power of growing out, either to produce a radical
hair (Bangid), or a new ramifying and fructiferous fila-
ment (Achlya).

Even mucilage, mixed with various other contents,
which had escaped from injured Algae-cells into water,
and lay scattered in detached portions in this, sometimes
presented to me a bounding layer on the surface, which I
could not distinguish from the delicate membrane, such as
occurs on young cells. This bounding layer is wanting
to mucilage which has just escaped from the injured cell.
It is evident that it was produced by the mucilage.

These three facts prove, that organic, unazotized mole-
cules are secreted on the surface of living vegetable mucilage,
and these inclose the mucilage in the form of a membranous
layer* Whether mucilage be free or lie in contact with

* I here presume to add a remark on the term " mucilage" (schleim,
mucus). Most vegetable physiologists use the word, partly for vegetable
gelatine, partly for gum ; on the other hand, Schleiden applies it to the

CELL-FORMATION. 125

a membrane makes no difference as to its function. Even
in regard to the secreted layer, the distinction is not
essential, and exists only in so far that in the one case
this is called a membrane, in the other a layer of thicken-
ing. That there is no distinction between membrane and
thickening layer is proved by the formation of membrane
on the surface of a partially free mucilaginous layer, in
the cells of Algae. There the newly-formed piece of mem-
brane passes continuously into the innermost layer of
thickening ; or, in other words, the gelatinous deposit
secreted simultaneously over the whole surface of the
mucilaginous layer, appears in some places as a layer of
thickening, in others as a membrane.

If we conclude from the fact that the mucilage pro-
duces membrane or layers of thickening through secretion
on its external surface, and that there is no essential dis-
tinction between them from the formation of membrane
to that of cells, it follows logically that the membrane must
originate through secretion from the mucilage in all cell-
formation. This conclusion is very strongly supported
by certain phenomena in cell-formation itself.

proteine-compounds. An agreement between these views would not be
readily effected. In addition there is a substance which is called schleim by
both parties ; this is the homogeneous, thickish, colourless matter, forming
the total contents of the young cell, and the homogeneous, denseish, colourless
ingredient in the contents of old cells. If, now, schleim is to have the sig-
nification of one or other (ternary or quaternary) of the chemical substances,
it is incorrect to call the last-mentioned schleim. Eor this is certainly not a
pure substance, but at the least a mixture of ternary (gum and sugar) and
quaternary substances (proteine-compounds). This mixed matter is of the
greatest importance in the life of the cell, since it is effective in cell-forma-
tion, in the first instance fills the whole cavity of the cell, subsequently lies
upon the wall as mucilaginous layer, and traverses the cavity in currents
of circulation, lastly, forms the contents of many utricles. (See the
Essay ' On the Utricular Structures' in this volume.) It therefore requires
a special name, and since the term schleim, on the one side, may certainly be
spared from the nomenclature of ternary and quaternary organic matters,
while on the other it is already in use in the proposed sense, it would be
most advantageous to name the homogeneous denseish substance of the
vegetable cell-contents, composed of mingled ternary and quaternary matters
schleim. [Mohl has proposed, and Schleiden and others have adopted, a
better name, protoplasma, which has solely a physiological value, confusion
resulting from the misuse of a term having a chemical signification. A. H.]

1:20 VEGETABLE CELLS.

I have already remarked* that, in the abnormal cell-
formation in the cells of Alga?, a great multiplicity of
conditions occur in regard to the contents which produce
the membrane, and that a continuous series of gradations
exists from abnormal cell-formation to the normal phe-
nomena of cell-life. Sometimes a free cell is formed by
a small portion of homogeneous mucilage, sometimes by
a larger quantity of mucilage containing chlorophyll and
starch ; sometimes a considerable portion of the cell-
contents produce, here a membrane, there layers of thick-
ening ; sometimes the whole contents form either a
perfect membrane, or partly membrane, partly thickening
layer, or a complete layer of thickening. These various
phenomena are all connected by a number of intermediate
states, so that the necessary cause and nature must be
alike in all.

Since now, in some cases (if, namely, the mucilage
borders on an existing membrane or on water) it is cer-
tain that the membrane cannot originate in any other
way but through secretion from the mucilage, which is
afterwards inclosed, we must undoubtedly assume a like
origin in the other cases, where a part or the whole of the
surface of the mucilage is bounded by fluid, unazotized
or azotized contents. Consequently, in free abnormal
cell-formation, the membrane is formed through secretion
from the portion of contents which becomes a cell, and
by no means in any other way from the rest of the
contents of the parent-cell.

Passing from the abnormal parietal and free cell-
formation to the normal parietal cell-formation, we find
here also that the phenomena themselves lead to that
assumption through a conclusion deduced from analogy.
I have already discussed this at length in the section on
parietal cell-formation. The contents of the parent-cell
divide into one or more portions. The new membrane
originates in some places between the mucilaginous con-

* Pages 97, 99, 114.

CELL-FORM ATION. 127

tents and the wall of the parent-cell, in some places be-
tween the mucilaginous contents and the wall of the
simultaneously-produced fellow secondary cell. Here
again no other hypothesis is possible but that of the
origin of the membrane through secretion from the muci-
laginous contents. The surface of the contents secretes
layers of thickening up to the moment when cell-forma-
tion begins. The layer secreted at this moment is the
rudiment of the membrane of the new cell.

In free normal cell-formation, lastly, the phenomena,
if they do not exclude every other theory, are not un-
favorable to the hypothesis founded on analogy. The
most indubitable case is that in Achlya, where the spo-
rangium-cell sometimes originates by parietal, sometimes
by free cell-formation. Since here, in the one case (in
the parietal mode of origin) the membrane is produced
through secretion from the contents, it is certain that the
same happens in the other case. In the formation of the
germ-cells of Algae, Lichens, and Fungi, and in that of
the endosperm-cells in the embryo-sac, the phenomena
admit of a twofold explanation : either that the membrane
is a secretion from the portion of contents becoming
isolated, or that it is a deposit from the fluid of the cell
surrounding the isolated portion. Schleiden and Schwann
have, as is well known, maintained the latter explanation.
They found an analogy for it in the inorganic crystalliza-
tion, but I believe that there is no analogy to it in the
region of organic processes. On the other hand, the
former explanation finds its certain analogy in lignifica-
tion, in abnormal free and parietal, and in normal parietal
cell-formation. Thus there exists no reason why we
should not assume the origin of the membrane in conse-
quence of secretion from the contents, to be certainly
established also in free normal cell-formation.

The examples of the formation of membrane, in the
manner we have just seen, fall into two categories :
1, Those in which the phenomena directly require the
hypothesis that the membrane is formed through secre-

128 VEGETABLE CELLS.

tion from the contents ; 2, Those in which the phenomena
do not oppose this hypothesis, and at the same time do
not afford the slightest probability to any other assump-
tion. Thus no objection can be made against the con-
clusion from analogy, so much the less that the examples
of the second category do not include any single point
not found in the first category which might suggest a
process of a different nature.

From the foregoing discussions I must express the
definition of the formation of membrane in the following
terms : The cell-membrane is an investment lying upon
the surface of the contents > secreted by the contents them-
selves. The membrane of a cell is the product of its own
contents, as well in the beginning as subsequently. It
does not originate through the chemical action of one
substance upon another of different kind ; its formation
is an organic process, and in fact a process of secretion.
This theory of the formation of membrane is, as we have
seen, an immediate consequence of the facts, and avoids
those formal and material errors which are connected with
the theories of Schwann and Schleiden, and of which I
have spoken elsewhere in the definition of a cell.

The other essential epoch in cell-formation is the in-
dividuali^ation of a portion of the contents, which becomes
transformed into a new cell. Here again we must, at the
outset, distinguish between normal and abnormal cell-
formation. In abnormal cell-formation, the mode in
which the contents become individualized does not de-
pend on a definite law, but on mere external, accidental
circumstances. Large or small, many or few, portions
of contents become isolated and form new cells. The
idea of individualization of portions of the contents is
here altogether vague, since it gradually loses itself in the
opposite process.

I have already repeatedly noticed a series of phenomena
which the abnormal cell-formation presents in living cells.
The one extreme is exhibited where a minute free portion
of contents, composed of mucilage alone, becomes clothed

CELL-FORMATION. 129

by a membrane ; the other is where the contents of a
cell become detached from the walls only in places, and
produce new membrane on these places. One extreme
is a perfect cell-formation ; the other merely a slight
alteration of the form of the cell, combined with a partial
production of new membrane. One extreme is the pro-
duction of a new individual cell ; the other is the re-
organization of the partially injured individuality.

Between the two extremes occur a quantity of inter-
mediate stages, in which it is doubtful, in particular
cases, whether the old cell persists in an altered form, or
whether a new cell has replaced it. The fact is certain,
but the explanation is furnished by the observer. In all
those cases where the mucilaginous layer is merely de-
tached in places from the cell-wall, and produces pieces
of membrane, we always recognize the cell as the same,
with partially altered form and membrane. In all those
where portions of the contents become isolated, and
complete their separation from one another by the forma-
tion of membrane, we suppose new cells to take the place
of the parent-cell ; but if the entire contents of a cell are
detached from the wall, become isolated, and then form
a new complete membrane, we may call this a cell-forma-
tion, as well as we can call it formation of membrane.
The contents of the old and new cell are exactly the same,
the walls altogether different. Is a new individual
formed, or is the individual merely reorganized (through
regeneration of an organ) ? This case stands just mid-
way, and admits either explanation equally well. A step
toward one side (if not the whole contents, but merely a
portion, become detached from the wall), or one step
toward the other (if not the whole contents, but merely
a portion of them become completely isolated), will strictly
decide the explanation to be either reorganization of the
cell, or cell-formation.

The transition occurs again in a different way. If the
contents of a cell become detached from the wall in one
place, and acquire a new coat of membrane over this

130 VEGETABLE CELLS.

place, while a larger or smaller portion of the contents
are lost and become dissolved, through the separation, we
shall always regard a cell altered in this manner as the
same. It has indeed lost a part of its contents, and
acquired a partially different membrane ; but the greater
portion of its contents and membrane remain unaltered.
On the other hand, if a very large quantity of the contents
separate and disappear by solution, we may regard the
remaining portion, which becomes inclosed by a mem-
brane, as we please, either as the altered original cell, or
as a new one. If the detached portion of contents, instead
of undergoing solution, remain living and acquire a mem-
branous coat, two new separate cells will have replaced
the original cell. We must, therefore, assume that cell-
formation takes place here. Of the three cases mentioned,
the first is undoubtedly a reorganization of the individual
cell ; the third, undoubtedly a production of new cells ;
the second is either one or the other, according as it is
compared with the one or other process.

The result of these considerations is : that abnormal
cell-formation cannot establish any firm and absolute defi-
nition for the process of individualization of the portions
of contents for the purpose of cell-formation ; because
this process is essentially uncertain, and merely different
in degree from that which stands in opposition to it.*

Passing now to normal cell-formation, we find the pro-
cess of individualization of the portions of contents con-
nected with determinate laws. We find no phenomena
here, forming transitional stages towards a different pro-
cess, in any way, towards that of mere reorganization of
the cell. The individualization of the contents for the
purpose of cell-formation appears, in general, under four
different forms :

1. Solitary ', minute portions of the contents become iso-

* I was compelled to establish this fact somewhat at length, because
many abnormal phenomena of cell-life have already been taken as evidence in
discussions on cell-formation. They are testimony for the formation of mem-
brane, but not for that of the cell generally.

CELL-FORMATION. 131

lated in the interior of the rest of the contents of the
parent-cell (Formation of free germ-cells in Algae, Lichens,
Fungi, and of the endosperm-cells of Phanerogamia.)

2. The entire contents of one cell, or of two cells con-
nected by conjugation, unite into one free, globular, or
ellipsoidal mass (Formation of the germ-cells of Zygne-
macese).

3. The entire contents of a cell divide into two or more
portions (parietal cell-formation in cell- division.)

4. The entire contents of a short branch of a cell, or
of the terminal portion of a longer branch, separate from
the rest of the contents of the cell (parietal cell-formation
in the so-called constriction, as, for instance, in the for-
mation of the germ-cells of several Algae \Vancheria, &c.]
and many Fungi).

The phenomena exhibited by the individualization of
the contents, in normal cell-formation, are externally the
same as in abnormal cell-formation. In abnormal cell-
formation, external, accidental influences exert a disturb-
ing action on the life of the cell. In normal cell-forma-
tion, the presence of a 'nucleus must be especially counted
among the conditioning causes. I have demonstrated it
to be in the highest degree probable that nuclei are always
present both in parietal and free normal cell-formation.
They are wanting in abnormal cell-formation, and in this
lies the essential distinction between normal and abnormal
cell-formation.

If we may presuppose the necessity of the presence of
a nucleus in the normal individualization of portions of
contents, the processes must be completed in the follow-
ing ways, in the four forms of individualization just
mentioned :

1 . One or more nuclei originate in the contents of the
cell. Each of these collects the contents in its immediate
vicinity upon its surface. These portions of contents are
composed of homogeneous mucilage, or mucilage with
which is intermingled other assimilated substances, such
as chlorophyll or other colouring matters, starch, oil, &c.

VEGETABLE CELLS.

Through this, free cells originate in the contents of the
parent-cell. The nuclei are in some cases already visible
during the individualization of the contents ; sometimes
they do not become visible until afterwards ; in certain
cases they have not hitherto been perceived.

2. A single nucleus originates, which lies in the midst
of the accumulation formed by the whole contents of one
cell, or of two conjugated cells. This accumulation forms
a single free cell in the empty cavity of the parent-cell.
The nucleus does not become visible until the develop-
ment of the germ-cells to new plants.

3. Several nuclei originate in the contents of the pa-
rent-cell, distributed in a regular arrangement. Each of
these individualizes the contents of the parent-cell, through
attraction, with a force only limited by that of the other
nuclei. At the limits which separate the field of action
of each nucleus, which, ceteris paribus, are equally distant
from each pair of nuclei, the membranes are formed. In
other words, the whole contents of the parent- cell divide
into just as many portions as there are nuclei ; a nucleus
lies pretty nearly in the middle of each portion ; the por-
tions are separated from each other by an extremely nar-
row space, in which each portion of contents secretes its
own membrane. The parent-cell divides, by parietal
cell-formation, into several secondary cells.

4. A single nucleus originates in a short branch (in
the prolonged part), or in the terminal portion of a longer
branch of a cell. Under its influence, the whole contents
of the short branch or of the terminal portion of the longer
one, separate from the rest of the contents of the cell
and form a new cell, the membrane of which is partly
parietal (in contact with the internal surface of the branch),
and partly free (directed towards the cavity of the older
cell). But the parent-cell, excepting in the loss it has
suffered, remains unaltered. The nucleus has never been
perceived yet during the actual process ; it not unfre-
quently becomes visible afterwards.

The definition of the INDIVIDUALIZATION OF THE CELL-

CELL-FORMATION. 133

CON TENTS /or the purpose of normal cell-formation is con-
sequently this ; that a nucleus is formed, and that this
nucleus individualizes a portion of the contents by attrac-
tion. The portions may be indeterminate or determinate.
If the former, the quantity of contents isolated around
the nucleus is uncertain. If the latter, either the whole
contents or a definite fraction (J, i, i, , |, i) becomes indi-
vidualized around the nucleus, the denominator of the
fraction giving the number of simultaneously individu-
alized portions of contents.

CELL-FORMATION is the individualization of a portion
of contents, immediately followed by the formation of
membrane. The definition of cell-formation depends on the
combination of the two constituent definitions. Normal
cell-formation, therefore, in so far as it is endogenous,
consists in this:

A nucleus is formed in the contents of the parent-cell ;
this individualizes a portion of the contents by attraction,
and this individualized portion becomes coated with a
membrane by secretion over the whole of its surface.

e/ t/ c/

But since, in animals, cell-formation also occurs outside
cells, likewise commencing with a nucleus, and since in
the generatio spontanea the first cells of plants or animals
are formed either in external media or in other cells, in
such a manner that these certainly cannot be regarded as
parent-cells in the proper sense of this word, the definition
of normal cell-formation must be brought into a more
general expression.

In organic substances (which in animal cell-formation
must contain proteine-compounds, in vegetable, azotized
and unazotized compounds, such as albumen, gum and
sugar) a nucleus originates which, by attraction, indivi-
dualizes a portion of the organic substances surrounding it;
the individualized portion of organic substances becomes
clothed by a membrane through secretion over its whole
surface.

In vegetable cell-formation the substance secreted as
membrane is composed of ternary unazotized substances.

134 VEGETABLE CELLS.

The homogeneous substance lying on the surface of the
cell-contents and secreting the membrane, is mucilage
(schleim) (albumen mixed with gum and sugar). Either
the whole portion of contents individualized through the
influence of the nucleus consists of mucilage, or of muci-
lage with which other matters are mingled ; in the latter
case, however, the extreme outer surface is always formed
by mucilage. There are two especial properties of the
mucilage which are of essential assistance in the formation
of cells. The first is this, that whenever it becomes free
or isolated, it acquires a perfectly smooth surface, on
which the strongest magnifying power fails to discover the
slightest prominences or depressions. A second property
is, that where mucilage and other solid matters unite into
a definite form, a portion of the mucilage always forms a
superficial layer, the other substances being as it were re-
pressed by it. The latter may be seen, both when the
contents of a ceh 1 (especially of the cells of Algae) flow out
in water, and when through injury to the cell the contents
become retracted from the walls, and divided into isolated
portions. Through the homogeneous mucilage forming a
continuous enveloping layer, or any portion of contents
becoming individualized, this becomes at once a con-
nected whole with definite boundaries. Through the
mucilage possessing a perfectly smooth external surface,
is produced a membrane, continuous and smooth, even
at the very first moment of its formation.

Having become acquainted with the definition of cell-
formation, and in it the conditions necessary to the pro-
cess, we will now examine more closely some of its
external relations. The cell may originate in another cell
or outside cells. In the vegetable kingdom the law holds
that all normal vegetative cell-formation takes place solely
inside cells ; moreover, that all normal reproductive cell-
formation, for the purpose of propagation, likewise occurs
exclusively in the interior of cells. Only the first cells
of individuals originating through (jeneratio aquivoca, are
formed outside cells.

CELL-FORMATION. 135

Schleiden was the first to express the formation of cells
in cells as the universal law for vegetables. In many
researches on the most diverse orders of the vegetable
kingdom, and on the most varied organs of plants, I have
never met with the formation of cells outside cells. Minute
investigation in the proper stages of development always
shows that the new cells proceed from parent-cells.

Meyen,* indeed, makes a portion of the cellular tissue
of the young anther become dissolved, and new cells to
originate in the homogeneous mucilage. But I have
shown that Meyen mistook a very thin, delicate tissue
for amorphous mucilage.f

Mirbel j considers that the cambium in the root of the
Date-palm consists of an amorphous, homogeneous, thick
fluid,in which the cells are formed. But in the roots of
the Date-palm, as in all other cambium in entire sections,
I find a continuous cellular tissue, and never any inter-
ruption by an amorphous mass.

Schleiden came to a similar result with regard to the
cell-formation in cambium.

Endlicher and Unger || recently affirm cells to originate
also through intercellular formation, appearing first as
cavities in the intercellular substance, and acquire proper
walls subsequently through the condensation of the latter.
But if cells lie in a gelatinous mass, this is by no means
a proof that they have originated in this. The history of
development must decide whether the cells have been
produced from the gelatinous mass, or the gelatine from
the cells. My researches on the Algae and Fungi show
to me that, without exception, the cells exist first, and
the gelatinous matter which subsequently surrounds them
is produced by the cells. In the simple plants, such as
Nostoc, Palmella, &c., the development may be traced
from the very first cell. In the more complicated plants,

* Physiologic, Band iii, 119 ; pi. xii, fig. 2.

Zur Eritwickelungsgesch. des Pollens, p. 10, figs. 2-7, 31-36, 47, 48.
\ Nouvelles notes sur le Cambium ; Archiv de Museum, tome i, pi. xxi.
Grundziige d. wiss Bot. 1st Ed. vol. i, p. 199.
j| Grundz. der Botanik, p. 33.

136 VEGETABLE CELLS.

as, for instance, the Fucoideae and Lichens, a thin-walled,
parenchymatous cellular tissue exists at first, and the
cells subsequently separate while gelatine makes its ap-
pearance between them. In many of the more complex
Algae and Florideae I have been able to trace the develop-
ment of the organs, step by step, from the firsit cell, and
to bring this within definite laws of cell-formation. It
thus becomes possible to determine in the perfect tissue,
in which parent-cell each cell has originated. But it
renders the hypothesis of an intercellular formation of
cells wholly impossible.

Quite recently, Mettenius* has stated that the cells in
the ovule and in the anthers of the Rhizocarpese, originate
in an amorphous fluid, which is inclosed in a sac formed
of a simple layer of cells. My own researches, however,
do not agree with this description. I have many times
seen a delicate- walled parenchyma during the cell-forma-
tion in these organs, and therefore am compelled to believe
that the origin of the parent-cells of the pollen in the
Rhizocarpese takes place in a similar way to that in the
Phanerogamia. In a single organ, namely, in the anthe-
ridia of Mosses and Ferns, I am still in doubt regarding
the cell-formation, not on account of anything unfavorable
to the theory, but because generally nothing at all is seen.
It is, besides, to be remarked, that the spermatic vesicles
which are formed there are without doubt, utricles ana-
logous to cell-nuclei (not to cells), and further, that on
the other antheridia (of the Characese and Florideae) no
extra-cellular cell -formation takes place.

The law being established, that in plants both the
vegetative and reproductive normal cell-formation takes
place only inside parent-cells, the relation of the cell-
formation to these parent-cells next demands consideration.
The first question is, in what number and what position
do these secondary-cells originate in the parent-cell.

The number of secondary cells is determinate or inde-

* Beitragc z. Kermtniss der Rhizocarpeen, p. 10 (1846).

CELL-FORMATION. 13?

terminate. In general, the rule may be stated, that in
vegetative cell-formation only two cells are formed from
one parent-cell, in reproductive cell-formation, on the other
hand, the number varies from one to an indefinite quantity,

i/ / _Z / '

and that here the smaller numbers (1, 2, 4, 6, 8) are con-
stant, while the larger numbers (5 to 100, or more,) vary.
In the first place, as to vegetative cell-formation, from
comprehensive researches in the Algae, Fungi, Lichens,
Florideae, Mosses, Charas, in the vascular Cryptogamia
and the Phanerogamia, I believe I am justified in ven-
turing to express, as an universal law, that here always two
secondary cells originate in one parent-cell, or, in other
words, that one cell divides into two. Views and repre-
sentations opposed to this I am compelled to regard as
certainly incorrect.*

In reproductive cell-formation, the cells originate in
various numbers in the parent-cell. One cell is found
constantly in the formation of the germ-cell of the Zyg-
nemaceae, and several other Algae and Fungi, also in the
formation of spores and pollen-cells in the special parent-
cells. Two, four, six, or eight cells are formed constantly
in one cell in the formation of the germ-cells of several
Algae, many Fungi, of Lichens, also in the formation of
the special-parent-cells of the Florideae, Hepaticae, Mosses,
Ferns, and Lycopodiaceae. The number varies from two
and four to eight in the formation of the special-parent-
cells of Phanerogamia. The secondary-cells originate in
indefinite quantity in the formation of the germ-cells of
many Algae and Fungi, and in the formation of the endo-
sperm-cells in the embryo-sac.

The position of the secondary cells in the interior of

* Thus Schleiden and Vogel figure three young cells in an epidermis-cell
which has grown out into a hair. (Beitrage z. Entw. der Bluthenth. bei d.
Legumin., Act. Acad. C. L. C. N. C. xix, 1, pi. x, fig. 38.) Schleiden also
represents three secondary cells in a parent-cell from the terminal shoot.
(Beitr. z. Anat. d. Cacteen ; Mem. de 1'Ac. Imp. des Sc. de St. Petersbourg,
Ser. vi, t. iv, pi. viii, fig. 9.) In both the places named I have satisfied
myself that only two cells are formed in a cell ; that when, however, one of
the latter quickly divides again, the appearance may readily be mistaken for
three cells originating in one parent-cell. See pp. 119, 120.

138 VEGETABLE CELLS.

the parent-cells is partly regular, partly irregular. The
arrangement of the secondary cells is regulated by definite
laws in vegetative cell-formation, and these produce a
definite internal structure and external form of the organ,
as 1 have pointed out in another essay,* in Delesseria
Hypoglossum, and in the Mosses. In reproductive cell-
formation we usually find a regular position connected
with a definite number, an irregular arrangement with an
indefinite number ; when the germ-cells (in Fungi and
Lichens) originate in the constant number of 2, 4, 6, or 8
in one parent-cell, they mostly lie in a row in the axis of
the parent-cell, or else they exhibit a mode of arrange-
ment having a definite relation to this axis. When the
special parent-cells are formed in the constant number of
four together, they are arranged tetrahedrally about the
centre of the parent-cell. But when the cells originate
in indefinite numbers (as in the formation of the germ-
cells of many Algae and Fungi, and of the endosperm-
cells), there is nothing determinate in their relative
positions in the parent-cell.

After the number and arrangement of the secondary
cells within the parent-cells, comes the further question,
in what way do the particular parts of the parent-cell co-
operate in cell-formation? The membrane, as we have
seen, in free and parietal cell-formation, bears no imme-
diate share in the production of the new cell. It is
merely the contents of the parent-ceh 1 which here come
into consideration. Normal cell-formation may, in re-
ference to the material relation of the parent-cell and
secondary cell, be brought into the following categories :

1 . A secondary -cell originates in a parent-cell, entirely
filling up the cavity of the latter, and perhaps including
the whole of its contents. Under this head are found
the origin of the spore-cells in the special parent-cells of
the four-spored Cryptogamia, namely the Floridese, Hepa-
ticse, Mosses, Ferns, and Lycopodiacese, and of the pollen-

* Zcitsch. fur wiss. Botanik. Schleiden und Nageli. Part ii (1845).

CELL-FORMATION. 139

cells in the special parent-cells of the Rhizocarpeae and
Phanerogamia. The special parent-cells contain a central
nucleus. This disappears, and soon afterwards we find
in the special-parent-cell a spore- or pollen-cell completely
filling it, with a (primary) lateral nucleus. How this cell
is formed, whether free in the contents, or around the
whole contents, is still unknown.

2 . The cen tral primary nucleus of the parent-cell divides
into two secondary nuclei; the entire contents separate
into two portions, each of which has one of these nuclei
in its interior. The parent-cell divides by parietal cell-
formation into two cells ivith central nuclei. This process
is met with in the formation of cellular tissue in the
Algae (and the Lichens ?), and in the reproductive cell-
formation of many one-celled or several-celled genera of
Algae.

3. A central nucleus appears in the parent-cell, and di-
vides into two secondary nuclei. The contents separate
into two portions, each of which includes one of these two
nuclei. The parent-cell divides, by parietal cell-formation,
into two cells with lateral nuclei. Here are to be enume-
rated the formation of cellular tissue of (most Fungi ?)
Florideae, Hepaticae, Mosses, Ferns, Lycopodiaceae, Cha-
raceae, Equisetaceae, Rhizocarpeae, and Phanerogamia. It
is still uncertain whether the central, dividing nucleus is
the primary nucleus of the parent-cell, becoming detached
from the wall, and advancing into the centre, or whether
it is a newly-formed nucleus, originating in the centre,
after the solution of the primary, lateral nucleus. The
settlement of this question will at once decide, whether
this cell-formation is actually different from that mentioned
under 2, or is to be considered as merely a peculiar
modification of it. With regard to the formation of
the stomates of the Phanerogamia, I have expressed the
opinion, that the primary nucleus of the epidermal cell
becomes absorbed and that a new central nucleus is
formed.* Mohl,f on the other hand, asserts of the same

* Linmca, 18 2, p. 238. f Vcrmischte Scliriften, p. 258.

140 VEGETABLE CELLS.

case, that the primary nucleus does not become dissolved,
bat that it is this which becomes divided.

4. Four nuclei are produced, probably by the division
of the primary central nucleus into two, each of which
becomes again divided. The four nuclei assume a tetra-
hedral arrangement. The contents separate into four por-
tions, each of which contains one of those nuclei in its
centre. The parent-cell divides by parietal cell-formation
into four tetraJtedr ally -arranged secondary cells. I have
hitherto only observed this process in the new single-
celled genus of Algae Tetrachococcus*

5. The lateral (primary] nucleus of the parent-cell be-
comes absorbed. A new secondary nucleus makes its ap-
pearance in the centre of the parent-cell. At its sides
originate two smaller nuclei.

A. The secondary nucleus of the parent-cell becomes ab-
sorbed. The contents separate into two portions, each of
which contains one of those smaller nuclei in the middle.
The parent-cell divides by parietal cell-formation into two
cells with central nuclei. Each of the two nuclei divides
into two nuclei, and each of the two cells divides again
into two secondary cells by parietal cell-formation.

B. The two small nuclei divide, so that now four still
smaller nuclei lie around the large secondary nucleus in
the parent-cell.

a. The secondary nucleus of the parent-cell becomes
absorbed. The four small nuclei assume a tetrahedral ar-
rangement. The contents separate into four portions, each
having one of these four nuclei in the centre. The parent-
cell divides by parietal cell-formation, into four tetrahe-
dr ally -arranged cells with central nuclei.

b. Some, or all of the four small nuclei divide, so that
then from five to eight still smaller nuclei lie around the
large central secondary nucleus of the parent-cell. The
latter becomes absorbed. The contents separate into just

JL J

as many portions as there have been small nuclei produced,

* Nageli, Neuern Algensysteme. Zurich, 1 847.

CELL-FORMATION. 141

each containing one of them inclosed in its centre. The
portions of contents are symmetrically arranged around
the centre. The parent-cell divides by parietal cell-for-
mation into from jive to eight cells with the central nuclei.
The process described occurs only in the formation of
the special parent-cells in the sporangium of the four-
spored Cryptogamia, and in anthers. The modification
given under A, I have observed in Florideae, Mosses, and
Phanerogamia ; the modification B#, in Florideae, Mosses,
Ferns, Lycopodiacea3, Rhizocarpeae, and Phanerogamia;
the modification B, in Phanerogamia.

6. In one parent-cell, or in two connected by conjuga-
tion, the whole of the contents unite into a globular or
ellipsoidal mass, which is transformed into a free cell by
the formation of a membrane on its surface. Here refers
the reproductive cell-formation of the Zygnemacese. Most
probably, a central nucleus lies inside the contents, this,
however, is in any case different from the primary nu-
cleus of the one, or those of the pair of cells.

7. New nuclei originate in a parent-cell, its primary
nucleus becoming dissolved ; each of these collects generally
a very small portion of contents around it, which become
coated by a membrane. The cells originate by free cell-
formation in the contents of the parent-cell. Here be-
long the formation of the germ-cells in some Algae, in
many Fungi, in the Lichens, and the formation of the
endosperm-cells in the Phanerogamia.

8. A cell grows out into a branch, and divides by pa-
rietal cell-formation into two cells, in such a way that one
corresponds to the original cavity of the cell, the other to
the expanded part. Here are to be enumerated the for-
mation of branches in Algae, Fungi, Floridese, &c. It
exists probably in all plants, but may be recognised best
in the organs composed of rows of cells. The origin of
the cell-nuclei, and their participation in the cell-forma-
tion, is not yet certainly made out. It seems to me that
the two following modifications exist, in respect to this :

A. The expanding cell possesses a central (primary)

142 VEGETABLE CELLS.

nucleus before its growth begins, and retains it subse-
quently. A free nucleus also exists afterwards in the
prolonged portion, and this probably originates sponta-
neously and independently of the primary nucleus of the
original cell. Each of the two secondary-cells possesses
a central nucleus. This branching cell-formation holds
good for those plants in which the formation of cellular
tissue follows the rule described under 2.

B. The expanding cell originally possesses a lateral
(primary) nucleus. A second nucleus is subsequently
found in the prolonged portion, probably originating
spontaneously and independently of the primary nucleus.
Each of the two secondary-cells possesses a lateral nucleus.
This branching cell-formation occurs in such plants as
have their cellular tissue formed according to the rule
given under 3.

9. A cell grows out into a branch. The whole of the
contents of the branch (Jf it be short), or the terminal por-
tion of its contents (when the branch is long), becomes
isolated, and, by the formation of a membrane over its
whole surface, is converted into a cell, which corresponds
exactly to the cavity of the very short branch, or to the
whole of the end of a longer branch. In this way the
germ-cells of several Algse (e. g. Vaucheria) and Fungi
are formed. The behaviour of the nucleus is not yet
sufficiently known. The parent-cell possesses a lateral or
central nucleus ; the secondary cell likewise. The nu-
cleus of the secondary cell probably originates in the
branch, spontaneously and independently of the (primary)
nucleus of the parent-cell. This mode of cell-formation
is closely allied to that mentioned under 8. But there
the division always occurs by means of a septum, which,
as soon as it is thick enough, appears to be formed of
two layers of equal thickness. From this condition of
the wall we may conclude that two cells are simulta-
neously formed, dividing in the contents and cavity of
the parent-cell. Here, on the other hand, only one la-
mella can be perceived in the curved septum, which

GROWTH OF CELLS. 143

always has its convex surface directed toward the original
cavity of the cell, and this lamella belongs to the cell
formed in the branch. The cavity of the parent-cell at
first appears to be closed in merely by the secondary-cell,
subsequently it becomes closed in by its own piece of
membrane. In this kind of cell-formation, therefore, as
I believe, not two, but only one cell originates in the first
instance.

VI. -THE GROWTH OF CELLS.

The cell is very varied in form at the moment of its
origin. Cells produced by free cell-formation are always
globular or ellipsoidal. Cells originating in numbers in
a parent-Bell, by parietal cell-formation, have that form
which is produced by the division of the parent-cell by
straight or curved surfaces. The variations of form under
which these newly-formed cells appear are innumerable.
The cell not unfrequently originates in a mathematically
regular form, such as a sphere, ellipsoid, hemisphere, a
quarter, segment, or section of a sphere, a cone, truncated
cone, cylinder, semi-cylinder, a quarter, section, or seg-
ment of a cylinder, tetrahedron, cube, table, prismatic
column, &c. &c. ; but in most cases the form of the nascent
cell is either intermediate between these mathematical
figures, or exhibits all possible irregular figures, with
plane (more rarely curved) surfaces. The faces and angles
of the nascent cell are either all curved or all straight, or
mixed. The number of faces by which the young cell is
bounded varies, according to my investigations, from 1 to
about 32, the number of plane angles from to about 90,
the number of solid angles from to about 60.

From these facts it is evident how little the existing
theory of cells is applicable, setting out from the spherical
form, and deducing all the others from this as subsequent.
In general I hold it as a rule that the cell retains the
shape in which it originates. The shape is certainly
altered in many cases ; but taking a number as the test,

144 VEGETABLE CELLS.

this is by no means the rule. When the cell does become
altered in shape, it much more frequently happens that
it passes from the polyhedral into the spherical, then
from the spherical into the polyhedral form. This last,
indeed, I believe only occurs in the endosperm-cells.

The growth of the cells is of two kinds. Either the
whole contents of the cell are simultaneously transformed,
and the entire membrane expands at once I will call this
universal growth (allseitiges Wachsthum}, or new contents
are continuously formed at one point of the surface of the
cell, and with this new membrane I call this apical
growth (Spitzenwachsthum).

In the growth of the cell it is necessary to distinguish
between the growth of the contents and that of the mem-
brane. The former appears always as causal and primary,
the latter as caused and secondary. I shall here speak
only of the growth of the membrane. This is a growth
in thickness, and a growth in surface. The latter comes
particularly into consideration in treating of the growth
of the cell. It is conceived as an expansion resulting
from the intussusception of organic molecules, and called
a nutrition of the membrane. Without entering into any
criticism of this theory, I will merely investigate, with
what phenomena the growth of the membrane in surface
is connected, in universal and apical growth of the cell.

When, in universal growth, the cell does not change
its form, the membrane must expand uniformly in all
parts. On the other hand, when the shape is altered,
some parts of the membrane must expand more than
others. The membrane consists of surfaces. In each of
these faces the growth of the membrane must be decom-
posed into two factors, the expansion in the two dimensions
of a plane. In each dimension we have three possibilities :
either the expansion is +#?, 0, or x\ either there is an
expansion or none ; or a diminution of the face of the
membrane in this dimension. The last case, though rare,
does occur in a few instances. The growth of the surface
of a membrane, therefore, when the three possibilities in

GROWTH OF CELLS. 145

each dimension are combined, exhibits, in general, nine
possible cases, leaving out of the question that a multitude
of differences in quantity may occur. In a cell where
the growth is not uniform, we must distinguish at least
between two faces ; but cells occur with ten, twenty, and
thirty faces. Prom this it may be conceived what an infinity
of theoretically possible diversities exist for the universal
growth of a cell-mernbrane in its totality, since this pro-
ceeds from the combination of so many and, more than
this, such variable factors. I think, therefore, that in
particular cases the growth of the membrane may be
decomposed into its factors, but that it is impossible to
establish general rules.

Schleiden* considers that the variations of form of
cells result from unequal nutrition of the membrane, this
taking place only in those places where one cell is in con-
tact with another, or with a fluid ; moreover is more
vigorous in those situations where a more considerable
interchange of matters with other cells is going on, thus
at the ends of elongated cells more strongly than at their
lateral faces, f

Schleiden here starts from the facts, that the cells of
the epidermis and of the septa in the air-canals are
flattened ; that in stellate and spongiform cells only the
rays are in contact with other cells ; that where a current
of sap passes through a tissue, elongated cells are formed.
The example cited, of Caulerpa, does not belong here,
but to a growth of quite different kind, to apical growth.

The facts brought forward show the incontestible rule,
if not absolute and unexceptional law ; but I draw quite
a different conclusion from them, namely, that the cell-
membrane grows least exactly where the greatest inter-
change of substances with external bodies takes place, and
that it expands most where it is least occupied in receiv-
ing and giving off substances.

I will first mention some negative reasons against

* Grundziige d. wis. Bot. } 3d ed. i, p. 211.
f L. c. 250.

146 VEGETABLE CELLS.

Schleiden's theory. If this were correct, we ought con-
stantly to find the epidermal cell next the air flat ; but
there exist such as are not merely not flat, but of which
the radial diameter is considerably greater than the taii-
gental. Moreover, there ought to.be a thorough distinc-
tion of form between epidermal cells in contact with water
and those exposed to air, which is not the case. Again,
the epidermal cells next the air should have a flattened
outer surface, while in many cases they are elevated like
papillae. The cells of cellular plates (simple layers of
cells) which exist in air, for instance the leaves of most
Mosses, should be plane and flattened on both sides, which
they often are not ; while, on the contrary, cellular plates
lying in water, e. g. Algae and Moridese not unfrequently
have tolerably plane and flattened cells.

The positive reasons are more important than these
negative objections. Assuming that a cell with equal
diameters and cubical form becomes tabular, as may be
the case in many cells of the epidermis and of the septa
in the air-canals, the cell expands in two directions, which
I will call the tangental, while it expands little or not at
all in the radial direction ; but the two faces to which
the radial diameter is perpendicular, and which I will
call end-faces, then expand, although in contact with air,
not merely as much, but more than the other or lateral
faces. If we assume that all three of the diameters of
the cell were originally = 1, and such an increase to
occur that both the tangental diameters become = 3, the
radial diameter remaining = 1, we have the six faces,
each of which originally possesses a square content = 1 ,
now increased in such a way that the square content of
the four lateral faces = 3, while the square content of
each end-face has become = 9. In this case, therefore,
the superficial square expansion of a lateral face standing
in contact with other cells is three times, while that of an
end-face in contact with air is nine times the original
content. The latter is thus three times greater than the
former. It may indeed be objected here, that the

GROWTH OF CELLS. 147

expansion of the membrane depends solely on the nutrition
of the lateral faces, a part of the latter having become
end-face ; but for this hypothesis we require a displace-
ment of particular parts of the membrane, which has
never yet been demonstrated in the cells of plants. Be-
sides, I will show by another analogous example that
such displacement really does not exist.

According to Schleiden, the elongated cells owe their
shape to a current of sap, their ends being thus more nou-
rished. On the other side it is to be objected, that if a
cylindrical or prismatic cell is more nourished at its end
than at its lateral faces, the cell will become proportionately
broader not proportionately longer, since the ends deter-
mine the breadth of the cylinder or prism. But, according
to the possibility mentioned in the previous example, a por-
tion of the ends might continually become lateral faces ; so
that the ends would always remain the same size, while the
lateral faces would grow by the addition of new portions
of membrane, especially at both extremities (that is, near
the two end faces). That this is not the case, however,
is proved by such cylindrical or prismatic cells as have
fixed points, on their lateral faces, from the conditions of
which we may judge of the expansion of the membrane.
The points of attachment of the branch cells are fixed
points of this kind, in plants and organs of very simple
structure. In Confervacese and Callithamniaceae the
branches often originate at a very early period ; they are
attached to the upper ends of the cylindrical lateral faces.
Now, I observed in several cases that the superficial con-
tents of the lateral faces expanded ten to twenty times
more than the end faces, after the origin of the branch,
and that during this the branch always either remained
near the end, or was but slightly removed from it. The
lateral face is free,, and in contact with water ; the end is
in contact with a cell. According to Schleiden's theory,
the end face ought to be more nourished than the lateral,
since it effects the exchange of organic substances, while
the latter only serves for the absorption of inorganic

148 VEGETABLE CELLS.

nutriment. But just the reverse occurs here, and it is
actually the lateral face which expands. The assumption
that the lateral face is at all enlarged by portions of the
end face being pushed over to the side is impossible here,
since this must necessarily remove the branch-cell gra-
dually away from the end. That the branch-cell is not
all moveable on the lateral face, and that its situation
may be regarded as a safe criterion here, is proved by the
pore which the branch-cells of the Callithamniaceoe pos-
sess in the centre of their basal surface, and which cer-
tainly is not moveable.. The same may be observed in
Ceramiaceae and in other Floridese ; as, for instance, in
Polysiphonia, where the prismatic central cell frequently
bears a branch.

The objection that the plants mentioned live in water,
does not appear to me an important objection. For it
can certainly at most be asserted, that membranes in
contact with water will be more readily nourished than
those exposed to the air ; but not that membranes which
are moistened with water, possess a more active nutrition
than such as are applied against other cells. Besides
which, that objection will, in any case, not apply to Poly-
siphonia, where the elongated central cells, the growth
of which furnishes the same result, are wholly surrounded
by cells. In order further to settle that doubt, I may
add that cylindrical cells, the lateral faces of which are
exposed to the air and the ends applied to cells, also, as
for instance in several filamentous Fungi and many hairs
of Phanerogainia, exhibit a similar condition to that of
the cells forming the joints of the Callithamniacese ; that,
namely, their lateral faces expand considerably more than
their end faces, which here alone officiate in the recep-
tion and transmission of fluid nutriment. The growth
of the cell may be judged of here again by the relative
position of the branch-cell, and in addition by the rela-
tive position of the nucleus of the cell. Hairs consisting
of a single cell, are originally little hemispherical or
conical cells ; the parietal nucleus lies at some point of

GROWTH OF CELLS. 149

their periphery. The cell often becomes twenty to forty
times longer than it was at first. The cell is exposed
by all its surface, except the base, to the air ; it acquires
its fluid nutriment solely through the basilar surface.
If the membrane were principally nourished by the pas-
sage of nutrient matter through it, the basilar surface
alone would be much expanded. Now, since the nucleus
exists in the earliest condition, and since normally it is
not moveable on account of its lateral attachment, it
must, if the theory were correct, lie normally near the
apex of the enlarged cylindrical cell, from the growth of
the membrane at the base. But it lies, without distinc-
tion, sometimes at the apex, sometimes in the middle, and
sometimes at the base of the cell.

Observations on elongated cells, or vessels with spiral,
annular, or porous lignification, afford me similar results.
If such cells or vessels grow after the lignification has
become evident, the spiral fibres, the rings, or pores,
separate uniformly on the whole of the lateral surfaces ;
the best proof that such elongated cells or vessels expand
at their lateral faces, and not at their terminal faces,
where the current of sap passes through the membrane.

The question still remains of the behaviour of the
stellate or spongiform cells. They are at first parenchy-
matous, but by the secretion of air become separated
from each other in the greatest part of the surface, re-
maining united together only in places. At the points
of contact the cells elongate radially. Schleiden calls
this a " growing-out " (Auswachseri) . I see nothing in
it but a mechanical drawing-out of those places which
cannot separate from the other cells, through the air
which has been excreted. The growth of the cells is a
different and much more independent process, while the
radiating elongation of the stellate and spongiform cells
stands in direct relation to the quantity of air secreted.
The undulated and dentate epidermal cells, however,
which are dove-tailed into one another, and which
Schleiden also cites, prove in any case nothing for his

150 VEGETABLE CELLS.

theory, because a membranous surface, which is uniformly
bounded at all points by another surface of membrane,
must exhibit like behaviour at all points in reference to
nutrition and expansion.

From the facts here brought forward, I deduce the
conclusion, that the cell-membrane normally expands less
in those places ivhere it serves for the reception and
emission of the nutrient matters, or where it officiates in
an interchange of substances or a current of sap ; that,
on the other hand, it normally expands more at those
places where it has little or no nutrient matter to manage.

The apical growth is altogether different from the
universal growth of cell-membrane. In an earlier me-
moir on Caulerpa* I have pointed out this apical growth,
and discussed its characters at length.f The apical
growths may be observed most easily in the filiform,
branched, single-celled Algae and Fungi, as in Caulerpa,
Bryopsis, Achlya, &c. The lower part of the cell is
quite perfect ; little or no change occurs in the contents ;
the cavity is filled with clear, colourless fluid, and on the
wall lies the mucilaginous layer, with the chlorophyll-
and starch-globules : the membrane expands little or no
more. Toward the extremity the membrane becomes
thinner ; it is there that new contents and new membrane
are formed.

Immediately below the growing point lies a layer
or small disc of homogeneous mucilage. Below, this
passes into granular mucilage. To this follows a mass of
granular mucilage, in which originate starch, chlorophyll,
or other colouring matters. Still lower down, the solid
contents are deposited upon the walls.

* Schleiden und Nageli, Zeitsclir. f. wiss. Bot., Heft i, p. 134.

f Schleiden (Grundz. d. w. Bot. 3d ed. i, p. 212) confounds the apical
growth with the unilateral expansion of universal growth. It is by no means
clear to me how he imagines Caulerpa to be " a body which at least appears
like a single cell;" further, how " the most decisive proof" of anything can
be deduced from this appearance. That Caulerpa actually is a single cell,
follows, in my opinion, both from the researches I have published and from
its relationship to Bryopsis } Valonia, Codium, Halymeda, Udotea, Faucheria, &c.

GROWTH OF CELLS. ] 51

In the disc of homogeneous mucilage lying in the
punctum vegetationis, new homogeneous mucilage is un-
ceasingly produced, so that it remains the same, although
its lower edge is continually being converted into gra-
nules, and thus a portion unceasingly given over into the
zone of granular mucilage. The lower part of the latter
passes continually over to the coloured zone, since in it
the formation of colouring matters, starch-granules, &c.,
begins. The lower part of this continually applies itself
to the wall. In this manner the relations of the parti-
cular zones remain the same, although an unceasing pro-
cess of transformation is going on in them.

The apical growth is thus connected with a production
of contents, which takes place at the apex of the cellular
branch .

The membrane, which is of tolerably uniform thickness
in the lower part, becomes gradually thinner toward the
end of the cellular branch. Immediately over the disc of
homogeneous mucilage at the punctum vegetationis, the
membrane is very delicate, as in a young newly-formed
cell. It always remains so during the growth of the
branch. The gelatinous matter secreted by the disc of
mucilage is never appropriated to the thickening of the
membrane, but solely produces the expansion of it neces-
sitated by the growth of the apex. The membrane
expands likewise beneath the apex, and exhibits pheno-
mena there, similar to the universal growth of the cell
without apical growth ; at the same time it becomes
thickened by deposition of new layers.

The membrane therefore exhibits different behaviour
at the apex and beneath this part. At the apex it is
continuously nourished, so to speak, or, as seems to me
a more correct expression of the fact, new membrane is
incessantly produced ; the process of formation of mem-
brane going on there remains the same so long as the
apical growth lasts in unlimited axes, therefore, it is
unlimited also. Beneath the apex, the membrane grows

152 VEGETABLE CELLS.

by universal expansion and thickening up to a certain
point ; the process of formation of membrane then gra-
dually changes, and at length ceases.

The behaviour of the membrane at the apices of grow-
ing axes, and below these apices, exhibits exactly the
same opposed relation, as the formation of membrane in
cell-formation, and the universal growth of membrane
after cell-formation.

The apical growth, therefore, consists not merely of a
continuous production of contents, but also of a continuous
production of new menibrane at the apex of the axis.

Many cells with apical growth branch. They deve-
lope new branch-cells from their lateral surface, and
these elongate by a repetition of the apical growth. Cells,
also, which do not grow at the apex themselves, may send
out a branch which exhibits apical growth ; thus the cells
which grow out into an appendage, as in the cells consti-
tuting the articulations of the Confervas, to form a branch-
cell or a root ; many cells of young bark or epidermis, to
produce a hair or radical fibril; the sporangial cells of
Achlya, to discharge the motile cellules (pi. Ill, figs. 4,
6, 8), &c.

The phenomena occurring in these cases, when they
can be perfectly seen, as is possible, for instance, in
JSryopsis, are as follows : At some point on the surface
of the cell, where the branch is to be formed, a little disc
of homogeneous mucilage is produced immediately beneath
the membrane ; the mucilaginous layer here produces a
definite quantity of new mucilage. Then the cell- wall
becomes elevated into a little triangular, hemispherical, or
conical papilla, filled with homogeneous mucilage. The
papilla grows larger. The mucilage becomes granular at
the lower part. The granulation of the mucilage advances
continually upwards. The formation of chlorophyll, or
other colouring matter, and of starch, commences at the
base, and also progresses upwards. Then the mucilage
at the lower end begins to apply itself upon the walls,

GROWTH OF CELLS. 153

and the cavity is filled with colourless fluid, and comes
into communication with the transparent cavity of the
original cell.

The apical growth consequently commences with the new
formation of homogeneous mucilage, which produces new
membrane, and is continued by the persistence in new for-
mation of membrane-forming mucilage.

The question now comes, what is the relation of the
apical growth to universal growth and to cell-formation ?
The relation may be conceived in two ways : 1st, in
reference to the contents; 2dly, in reference to the mem-
brane. I will here, as in the universal growth, only enter
upon the latter point.

Schleiden and Schwann have strictly discriminated two
epochs in the development of the cell, namely, the origin
of the membrane, and its growth. This distinction, how-
ever, according to the view of cell-formation above estab-
lished, is certainly not so great as it appears in Schleiden 's
and in Schwann's theory. According to this theory, the
material for the original production of the membrane is
merely furnished by the mother-liquor, from which it
crystallizes ; the material for the growth of the membrane
is, on the contrary, partly or wholly furnished by the cell-
contents ; it is, namely, deposited from the contents, or
acquired by the endosmose and exosmose through the
membrane.

According to my theory of cell-formation, the original
production and the growth of the membrane have the
same material cause. For both purposes, organic matter
(in plants mostly gelatine) is secreted from the contents.
Origin and growth are, in the same way, a product of the
contents. They merely differ from each other relatively.
The original production of membrane gradually changes
into growth of membrane. A line of demarcation be-
tween them is quite arbitrary.

We see, therefore, that in this point the membrane
behaves just like every other organism. In none, cer-
tainly, does there exist a strict and absolute boundary

154 VEGETABLE CELLS.

between origin and growth. But if these two ideas are not
absolutely different from each other, we shall still let them
stand side by side as separate, in their relative difference.

When a cell is produced, a portion of organic matter
becomes individualized and inclosed by a membrane,
through secretion of organic substance. With CELL-
FORMATION, therefore, is connected an individualization of
organic matter and a formation of new membrane. When
apical growth begins at a point of the surface of a cell,
and a branch-cell is formed, a portion of homogeneous
mucilage appears, which secretes membrane from within
outward. With the commencement of APICAL GROWTH,
therefore, is connected the individualization of a portion
of the contents and a formation of neiv membrane, and,
with the continuous APICAL GROWTH, a continuous for-
mation of new membrane. I will consider these two
points a little more minutely.

When secondary cells originate in a parent-cell, definite
portions of the contents are repeated in such a manner,
that from that time they possess an individual existence,
having been up to that period undistinguished parts of
the parent individual. When a branch is produced upon
a cell, a definite portion of the contents is likewise sepa-
rated, and indeed a small portion of the mucilaginous
layer. This has from that time an individual vitality
wholly distinct from all the rest of the contents. This
little portion of mucilage is the rudiment from which a
new branch is produced. The branch of the ramified cell,
however, has quite the same import as the branch of a
many-celled plant. In the ramified cell, as in the many-
celled plant, it may, under certain circumstances, be de-
tached and become an independent, new, and individual
plant.

With the commencement of the APICAL GROWTH in the
formation of a branch, consequently, is connected the indi-
vidualization or separation of a portion of the contents, as
well as with cell-formation.

These secondary cells originate in a parent-cell ; the

GROWTH OF CELLS.

155

cell-membranes are produced around the individualized
portion by a new process of formation; the contents
secreting organic matter around. When a cell-branch
is produced, the individualized portion of contents forms
membrane only on its outer face. The growing out of
the cell- wall into a branch, however, does not, as I believe,
happen through unilateral nutrition, but by a new forma-
tion, especially from the fact that the wall of the parent-
cell or parent-branch is sometimes already tolerably thick
and lamellated before the branch is formed, and because
the growing out then assumes an aspect much more as if
the membrane of the parent-cell were pushed outward
and broken through, than as if it were expanded and
formed a branch through nutrition. The origin of a cell-
branch is distinguished from the origin of a cell by the
fact, that in the former membrane is formed only on one
side, in the latter over the whole surface ; and that in
the former the production of membrane is continued, in
the latter exists but at one short period. The APICAL
GROWTH, therefore, like CELL-FORMATION, is connected with
a formation of new membrane, but this is unilateral and
continuous.

In cell-formation, as soon as the membrane is formed,
it expands by universal growth. In apical growth, as the
new membrane is every moment produced, it also at once
begins to expand by universal growth. In both cases a
thickening of the membrane is combined, at the same
time, with the expansion. In both cases, moreover, the
expansion only persists for a time, and then ceases.
Finally, in both cases, the expansion of the membrane
is at all points equal or unequal; in uniform universal
growth, the cell or cell-branch is not altered in form ; in
irregular universal growth, the cell or cell-branch assumes
a shape different from that it originally possesses. As in
cell-formation, so also in apical growth, the new formation
of membrane and its growth, are only relatively different
from each other. Both are conditioned by the secretion

156 VEGETABLE CELLS.

of organic matters from the contents ; one passes into the
other without any definite boundary.

In reference to the formation of membrane, therefore,
we may establish the following distinctions : In CELL-
FORMATION, an universal and momentary production of
new membrane takes place. In APICAL GROWTH, an uni-
lateral and continuous formation of new membrane appears.
In UNIVERSAL GROWTH, wJdcli follows both cell-formation
and apical growth, an expansion and thickening of the
newly-formed membrane takes place.

The apical growth may begin in a cell in two ways, in
reference to its relations to the life of the cell in time
and space. In the first case, the apical growth begins
at once with the formation of the cell. A portion of the
contents becomes individualized, and secretes membrane
over its whole surface, whereby the cell is formed ; this
new formation of membrane continues at one point of the
surface, and the cell goes on growing in this direction.
The apical growth is here immediately connected with
the cell-formation. It begins directly after the origin of
the cell, before the universal growth has commenced.

In the second case, on the contrary, the cell first ex-
pands by universal growth. The membrane becomes
thickened; the contents are applied upon the internal
surface. Then a portion of homogeneous mucilage is formed
at one point of the surface, beneath the membrane, and
with this begins a new apical growth, as I have above de-
scribed it. The apical growth, therefore, does not here
follow immediately the cell-formation, but the universal
growth of the cell. It may occur, either in a cell with-
out apical growth, or in a cell with this, as on the lateral
part of a cell-branch growing at the apex, and always
appears as a growing-out of a point of the surface.

Through the first hind of apical growth the nascent
cell grows longitudinally. By the second kind the cell
branches.

The apical growth is also variable in its duration. It

GROWTH OF CELLS. 157

is LIMITED or UNLIMITED, according as it either ceases at
a definite time, or continues as long as the plant lives.
Apical growth is limited in all cells, growing at the apex,
of many-celled plants, also in many axes of unicellular
plants. It is unlimited in particular axes of unicellular
plants, as, for instance, in the main axis of Caulerpa,
Bryopsis, &c.

Apical growth is not found in all cells ; it is wanting
even in the great majority of cells. I have observed it
in unicellular Algae, as Caulerpa, Bryopsis, VaucJieria,
Falonia, Codium, Halymeda, Udotea ; in unicellular Fungi,
as Achlya, Bremia, Mucor, &c. ; in the terminal cells of
many Algae, Florideae, and Fungi, as, for instance, Con-
ferva, Dasydadus, Griffithsia, Polyactis, &c. ; in the roots
of many Algae, Florideae, and Fungi; in unicellular
hairs, or the terminal cells of many-celled hairs, in the
pollen-tube \ lastly, in all cells growing out into a branch,
as, for instance, in the cells forming the articulations of
many Algae, Florideae, Fungi, and the organs consisting
of series of cells in the higher plants, moreover in the
epidermal cells of the higher and lower plants, &c.

THE UTRICULAR STRUCTURES

IN THE

CONTENTS OF CELLS.

BY GAEL NAGELI.

TRANSLATED FROM

SCHLEIDEN U. NAGELl's ZEITSCHRIFT F. WISS. BOTANIK, 1846.
BY ARTHUR HENFREY, F.L.S.

UTRICULAR STRUCTURES.

IT is now some years since I first imagined that the
phenomena exhibited by various bodies occurring in the
cell-contents, and supposed to be solid, could only be
explained by the hypothesis that they were utricles, having
an inclosing membrane with inclosed contents. The
minute size of the objects in question, the uncertainty
which attributes to the appearances, on account of the dif-
ferent refractive power of different substances, and the cir-
cumstance that, already, repeated attempts to demonstrate
cellular or utricular structures in the cell-contents had
failed, impressed the necessity of great circumspection.
I now lay before the physiological public my researches
on the various forms of the contents in the various kinds
of cells, only after extensive observations, and a confir-
mation of them by a comprehensive critical inquiry.

Earlier and more recent theories, which declare the
cells to originate from starch- or chlorophyll-granules, and
from which might be deduced a cellular structure of
these granules, may conveniently be passed over as un-
founded.

Earlier authors, likewise, who use granule and utricle
as synonymous terms, and imply no distinction between
the two expressions, merit no special mention.

More recently, granules and utricles have been placed
in opposition to each other. The first are considered
to be solid, the latter hollow. After this definition,
the opinion that cells inclose only granules gradually
gained the upper hand. Better instruments and more

162 UTRICULAR STRUCTURES.

accurate methods of investigation showed that what had
been considered hollow, was really solid.

It was further attempted to show, in some granules,
that their mode of origin was analogous to that of crystals
and different from that of cells. They were supposed
to be produced by a nucleus, through the deposition of
layers on the outer surface.

Particular theorists persisted in the view that the for-
mations in the contents were utricles, the analogy to a
cell floating in their minds. The empirical proof that
such an agreement existed was not given.

In the present condition of the researches, therefore,
the questions are : whether the so-called granules exhibit
analogy with the crystal or the cell ; whether they are solid
or hollow ; whether they are formed by the deposition of
layers around a nucleus or in some other way ?

The first question is to be solved by the decision of
the last two. The second is, of course, to be answered
with the statement., that as a rule the so-called granules
appear solid. But this does not establish anything in re-
gard to their nature, since utricles may become or appear
solid in various ways, while granules may at the same
time be hollow. The answer to the third question alone
possesses scientific interest, since it decides incontestably
as to the crystal-like or cell-like nature.

Thus the entire question turns upon this : Do the
structures in the cell-contents originate by stratified depo-
sition from without, and do they fulfil a vital process cor-
responding to this crystal-like mode of formation ? or, on
the other side : Do they originate in a similar manner
to the cell ; do they consist, like this, of a membrane and
contents inclosed by this membrane ; and do they exhibit,
in general^ in reference to the contents and membrane, ana-
logous alterations during their life, to those we are
acquainted with in the cells ?

Some time ago I pointed out two utricular structures
in the contents of cells, the nuclear utricle,* and peculiar

* SchleidenundNageli, Zeits. f. wiss. Bot,, i, p. 68; Ray Trans., 1845, p. 246.

UTRICULAR STRUCTURES. 163

utricles producing starch and chlorophyll, in Caulerpa*
As a criterion of their utricular nature, I mentioned that
they do not originate by deposition on the outside, that
they inclose, in a complete membrane, contents distinct
from this membrane, fluid, semifluid, or, sometimes, par-
tially granular, that they grow by expansion of their
membrane, and transform their contents, and that they
propagate in the same way as cells.

In their essential peculiarities, therefore, these utricles
present the character of cells ; and we shall consequently,
in the decision whether anything is an utricle or not, take
especially into account the phenomena which are analogous
to the phenomena of cell-life. On the other hand, we shall
not determine whether anything is to be considered as an
utricle by the circumstance of its being hollow or solid.
Now, as cells occur which appear solid, either because
their very delicate membrane is quite filled with contents,
or because they are completely lignified, so also do such
utricles present themselves. Moreover, hollow structures
occur in the cell-contents which are not utricles.

I must enter more minutely into the latter point,
because it has very often led to error. Homogeneous
mucilaginous contents frequently exhibit transparent, co-
lourless cavities. These are globular when they are iso-
lated, or at least are not very near together ; they become
parenchymatous when they are closely crowded. They
vary much in size ; sometimes a great number find place
in one cell, sometimes one, two, or three, occupy almost
the entire cavity.

The mucilage in which these cavities appear is either
of equal density throughout, or it is denser at the peri-
phery of the cavities, and forms as it were a membrane
around them. In such cases the cavities appear like
utricles. This utricular structure is the more deceptive
the more the density of the mucilage immediately on the
surface of the cavity differs from that of the rest of the
interior of the cell ; it is still more deceptive when these

* Loc. cit. p. 149.

164 UTR1CULAR STRUCTURES.

cavities are larger and exist in smaller numbers, and when
the mucilage generally is more diluted and transparent.
The cavities have, on the contrary, more the aspect of
real, mere cavities, when they are small and numerous,
and when the mucilage is dense and opaque.

These cavities contain water. They are seen in most
cells which are passing from the condition in which the
cavity is wholly filled with homogeneous mucilage, to
that in which the mucilage is deposited upon the wall as
a mucilaginous layer. They also occur, not unfrequently,
at a subsequent period, when the cells contain a homoge-
neous mucilage besides the solid matters ; I have seen
them in this late stage of the life of the cell, from the
cells of Conferva, Siphonece, and Hyphomycetes, upwards
to the parenchyma-cells of the Phanerogamia. They are
here a normal phenomenon of cell-life.

The following is the probable mode of origin of these
cavities. Larger or smaller quantities of water separate
from the mucilage, and from physical laws acquire a
globular form. From the loss of water, the mucilage
contracts and becomes more dense. If the drops of water
lie long unaltered in the mucilage, the layer of the latter
in contact with the water coagulates through its influence,
as is always more or less the case with mucilage or albu-
men in water. In this way originate in the cell-contents
the colourless pseudo-utricles possessing a special mem-
brane. In conformity with their origin we never see any
other contents but colourless fluid in them, and never
observe any alteration in their apparent membrane.

That this explanation of the normally occurring utri-
cular cavities is correct, is proved by the abnormal forma-
tion of similar cavities, either when mucilage or albumen
is mixed with water, or when we allow water to act endos-
motically upon a cell containing homogeneous mucilage.
In both cases, similar globular, colourless, sharply-defined
cavities are frequently produced, the mucilage at the same
time contracting and becoming more dense and opaque.

These cavities in the mucilage must by no means be

UTRICULAR STRUCTURES. 165

called utricles, because the utricles, according to their
definition, are to be described neither as hollow spaces,
nor as hollow spaces surrounded by a membrane. They
must at least possess a special membrane and contents
subject to special metamorphoses.

1. Nuclear Utricles, Nuclei.

That the nuclei are utricles I have pointed out in an-
other place.* I then brought forward the following
reasons in favour of this opinion :

1. When the nucleus admits of minute examination,
in respect of size and density, a membrane, and contents
distinct from this, may always be recognised. The mem-
brane is only overlooked where the nuclei are too small,
or where dense contents are inclosed in a delicate mem-
brane. In the latter case, it is sometimes possible to
render the membrane visible by the action of re-agents.

2. The membrane is different from the contents (by
no means the outermost layer of them) ; it is not coloured
by iodine and is composed of gelatine, while the con-
tents are usually coloured brown by iodine and consist of
mucilage.

3. The contents exhibit peculiar transformations, which
are analogous to those of the cell-contents.

4. The membrane is proper to the nucleus (by no
means a mere deposit from the cell-contents), as is proved
by the propagation of the nuclei, which exhibits the same
phenomena as the propagation of the cells.

I may add here that the membrane of the nucleus may
sometimes be recognised very distinctly, when it becomes
expanded through endosmose of water, f

Schleiden \ thinks " that the nuclei become hollow

* Schleiden und Nageli, Zeitsch. f. wiss. Bot., i, p. 68 ; Ray Trans. 1845,
p. 246.

+ See p. 110 of this volume, pi. ii, fig. 9, , k, I, m.

J Grundziige, second edition, i, p. 199. Schleiden reproaches me for an
unsettled terminology. I have called the nucleus an utricle (blascheri), and

166 UTR1CULAR STRUCTURES.

afterwards, since there is no trace of a membrane in the
young free cytoblasts, and the origin of these even appears
to contradict my view." If the origin of the nuclei could
be observed, this would afford the most decisive proof of
one or other of the theories. Schleiden indeed describes
the origin of the nucleus as a confluence or conglomera-
tion of mucilage-granules and nucleolar-globules. I cannot
agree at all in this opinion. The commencement of the
formation of the nucleus may be quite definitely distin-
guished, while it is yet little larger than the globules of
mucilage, and may be traced onwards uninterruptedly,
any deposition of mucilaginous granules upon it being
out of the question. On this point I refer to the preced-
ing treatise, where it treats of free cell-formation in the
embryo-sac.* The origin of the nuclei free in the cell-
contents consequently affords no evidence in favour of
the assumption, that they are solid granules without a
membrane.

Schleiden says moreover, that " in young free cytoblasts
no trace of a membrane is found." The youngest nuclei
consist of a homogeneous substance. They are either
more dense than the surrounding fluid, or, as is frequently
the case in young parenchyma-cells, they are less dense
than the surrounding mucilaginous contents, and appear
like hollow spaces in it. In this youngest condition
examination certainly reveals no membrane distinguish-
able from the contents ; but this is not a circumstance
which can be of weight in judgment of the nature of the
nucleus. If optical instruments never become improved

used nuclear utricle (kern-bldscheti}, as synonymous with nucleus. I still
know of no better terminology to substitute for it. On the one side, the
analogy with the other kinds of utricle requires the name "nuclear utricle ;"
on the other hand, custom of language, brevity of expression, and the con-
trast with nucleolus, leads to the use of the term " nucleus." The confusion
of language is always greatest when conceptions are in process of discrimi-
nation. Selecting from a hundred examples in botany, does not the same
author use spiral vessel and spiral-fibrous cell, or liber-fibre and liber-cell as
synonymous, although in regard to the last example, the \vord fibre already
represents a perfectly definite conception.
* See page 103 of this volume.

UTRICULAR STRUCTURES. 167

so far as to enable us to distinguish the organic molecules,
there will always be still entire stages, in the development
of membranes, in which it will be impossible to distinguish
them from the homogeneous contents lying close to them,
and refracting light in a similar manner, and in which
their presence or absence must be decided on upon other
grounds.

The matter stands in a similar position in regard to
cells. The relation of contents and membrane is evident
in fully-formed cells, and it may be carried back by a
conclusion from analogy to the young cells. The young
cells in the embryo-sac, if they possess homogeneous and
not granular contents, often do not allow of our perceiv-
ing the membrane for a long time. Free germ-cells
(spores) of Algae, Eungi, and Lichens, mostly attain a
considerable size (at least as considerable as that of the
nucleus mentioned by Schleiden), without the possibility
of seeing the least sign of membrane, and yet they are
young cells, to which no one will deny a membrane.*
Moreover, cells occur also, about the size of young nuclei,
on which no membrane is visible in their lifetime. Among
these are some species of Protococcus, of Palmella, and of
other Palmelleae.

If, then, it is impossible to distinguish the membrane
from the contents in young free ceils, or in small free
cells generally, on the other hand the membrane is usually
visible in parietal cells at the moment when the cell
originates, and in those very places where the apposition
of the secondary cells forms a septum. The same occurs
in nuclei which originate by the division of a parent-
nucleus. Here, too, as in the division of cells, a septum
is visible, formed by the two meeting membranes. f

The comparison of those free cells is especially neces-
sary here, in which membrane and contents cannot be
clearly distinguished even in the fully- developed con-

* See pages 95-6 of this volume.

f Nagcli on Cells, Part I, Hay Transl. 1845, pp. 231, 246.

168 UTRICULAR STRUCTURES.

dition. In them the septum is visible, as a line, only in
the moment of the division. If, therefore, analogy with
other cells does not afford evidence of the presence of a
membrane to these cells, we can only draw conclusions
in respect to it from the septum which is seen in the
propagation ; but when we can succeed in bringing two
such free cells together, in such a manner that, where in
contact, they are reciprocally somewhat flattened against
each other, the two membranes become visible at once,
as a septum, while at the remainder of the periphery, as
before the union was the case at all points, nothing can
be seen of the membrane. I have accomplished this in
Palmella, Protococcus, and Saccharomyces.

Nucleus and cell then exhibit, in reference to their
membrane and its relation to the contents, the same phe-
nomena. In small and young free individuals, the
membrane and contents are not distinguishable at first
sight ; they do not become so until they have undergone
further development. In individuals, however, which
originate through parietal formation, the membrane is
evident as a septum in the earliest condition. The sole
distinction, following from the nature of the case, which
prevails between nucleus and cell is, that there are far
more conditions of the nucleus than of the cell, in which
a distinction between membrane and contents is impos-
sible, since the phenomena are represented on a con-
siderably smaller scale in the nucleus than in the cell.

After disposing of the objections above mooted, I pro-
ceed to a short exposition of the vital phenomena of the
nuclear utricle.

The nucleus originates in two ways : either free in the
contents of a cell, or by the division of a parent-nucleus.
If the nucleus originates by division, the phenomena
presented are similar to those in the division of a cell.
A septum becomes visible in the parent nuclear utricle,
dividing it into two halves (as in Tradescantia] ;* soine-

* Niigeli on Cell-Formation. Ray Trans., 1845, p. 184, pi. vii, fig. 20.

TJTRICULAll STRUCTURES. 169

times a nucleolus is visible in both halves (as in Antho-
ceros)* Since the division of the nucleus exhibits
exactly similar phenomena to the division of cells, I con-
jecture that a parietal formation of the nuclear utricle
must be assumed in the former case like the parietal
cell-formation^ in the latter.

The process just described may perhaps be borne out
by the facts known regarding the division of the nucleus
in animal-cells, j A large nucleolus appears in the parent-
nucleus, and divides ; the two secondary nucleoli retreat
from each other, and the nuclear utricle divides into two
secondary nuclear utricles, in such a manner that each of
them incloses a nucleolus.

If we may draw conclusions as to the division of the
nucleus from the division of cells, where the appear-
ances are the same, the parietal formation of the nuclear
utricle depends on the following events : Two nucleoli
appear in the parent nuclear utricle ; the contents of this
separate into two portions, each of which incloses a nucle-
olus, and becomes clothed by a membrane ; the membrane
of each of the new nuclear utricles is in contact partly
with the wall of the parent-utricle, aud partly with that of
its fellow, whereby a septum is produced in the parent-
utricle.

When the nuclei originate free in the cell-contents, the
circumstances are probably the same as those occurring
in the formation of free cells. It appears, namely, that,
first, a nucleolus is formed ; that a layer of mucilage is
deposited around this, and that the membrane of the nuclear
utricle is produced upon the surface of the mucilage. At
least the phenomena which can be observed in the forma-
tion of free nuclei allow of this explanation, which is
rendered probable by the analogy to free-cell formation.
I refer the reader, on this point, to the article " Free
cell-formation" in the preceding essay.

* LOG. cit. pi. vi, fig. 38.

f Loc. cit. p. 284.

% Kolliker, Entwick. dcr Cephalopod., pi. vi, fig. 68.

Page 95 et seq.

170 UTRICULAR STRUCTURES.

When the nucleus originates free, it is in its earliest
condition globular; when it originates with a fellow-
nucleus in a parent-nucleus, it is originally hemispherical,
but then, separating from its fellow, it rapidly assumes a
globular shape.* The nucleus retains its spherical shape
if central in its cell ; it only becomes ellipsoidal when
about to propagate ; but if the nucleus occupies a lateral
position in its cell, it becomes flattened on one side. Seen
in front, it then appears circular or ellipsoidal, while
laterally it seems more or less compressed.

The growth of the nucleus is sometimes very slight ;
sometimes an expansion of from twice to ten times its
diameter takes place.

The membrane of the nucleus never acquires any
considerable thickness, and never exhibits lamellation or
lignification as in cells.

Young nuclei, as well when they have originated free
as by division, generally possess homogeneous, colourless
mucilaginous contents. These may be dense and opaque,
or almost transparent. Subsequently they become gra-
nular, and we may then distinguish a transparent fluid,
mucilage-granules, starch- granules, chlorophyll-granules,
and drops of oil. Sometimes the homogeneous and
granular mucilage becomes deposited in the form of a
layer on the inner surface of the membrane, so that the
rest of the cavity is filled merely with watery fluid.
Sometimes the nucleus contains amorphous colouring
matter, so that it appears homogeneously redf or green. J
Amorphous chlorophyll may also clothe the inner surface
of the membrane, merely in patches. For more minute
particulars, I refer to my earlier essay on this subject. ||

There are generally from one to three nucleoli in the
contents of the nucleus; sometimes their number rises
to five and six. These nucleoli appear to be rarely absent.
Probably they are always present, and merely invisible

* Mgeli on Cell-Formation; Ray Trans., 1845, p. 184, pi. vii, fig. 20.
f Loc. cit, 1845, pi. vi, fig. 26. Loc. cit, pi. vi, figs. 36-40.

Loc. cit. pi. vi, fig. 41. || Loc. cit. p. 219 et scq.

UTRICULAR STRUCTURES.

171

on account of the minute size, or the opacity of the
nucleus. It is, moreover, probable, when at a later
period several nucleoli are met with in one nucleus, that
one alone existed in the first instance, and that the rest
have originated subsequently. I have already expressed
my opinion on this point in the section on " Free cell-
formation" of the preceding essay.*

The nuclei terminate their existence either by solution,
in consequence of weakness resulting from age, or by
propagation. The mode in which nuclei propagate, has
already been described when treating of their origin.

2. Spermatic Utricles.

The spermatic utricles are those in which the spermatic
filaments originate. Hitherto they have been called cells
or cellules. I formerly thought that they might have
the import of nuclear utricles, because always one such
utricle appeared in a cell in the antheridia of the Cha-
raceae, and because a nucleolus always existed previously
in each utricle.

Without intending now to deny this analogy, I believe
that the spermatic utricles are not perfectly identical with
the nuclear utricles ; in the first place, because the pro-
duction of the spermatic filament is a characteristic of
which the nuclei are wholly devoid, and further, be-
cause in animals, and probably also in plants,! several
such utricles are sometimes formed in one cell.

It is no proof of the spermatic utricles being cells,
that in the Elorideae, Mosses, and Ferns, they are applied
closely together so as to form an apparent tissue ; since
it is quite as possible as not, that they are produced in
one large cell resembling an embryo-sac, or that the cells
in which they are formed become dissolved.

* Page 107.

f Nageli on the Propagation of the Ehizocarpeee ; Zeitschr. fur wiss.
Botanik, H. 3, p. 188 (184=6).

172 UTRICULAR STRUCTURES.

The mode of origin of the spermatic cellules is unknown,
as also of the propagation.

The shape of the spermatic utricles is globular or ellip-
soidal when they are free parenchymatous when they are
closely packed together.

The contents are originally homogeneous and muci-
laginous. In this may be perceived globular utricles
or solid corpuscles, probably analogues of the nucleoli
of the nuclear utricle. Sometimes the homogeneous
contents subsequently become finely granular. Chloro-
phyll-granules are occasionally formed in them. In the
Rhizocarpea3* they also contain starch-globules (?). At
a later epoch a spermatic filament, f composed of albu-
men, is produced in each of these utricles.

3. Nucleoli (Kernchen).

The nucleoli were formerly regarded as solid bodies.
I have already remarked upon this, some time ago, that
certain phenomena connect themselves readily with the
hypothesis that they are utricles. \ The question is now,
whether there are grounds for this assumption.

That the nucleoli are not mere aggregations of mu-
cilage, follows from the circumstance that they always
present themselves with a very definite border. More-
over, it is sometimes possible distinctly to perceive on
them a membrane different from the contents. The con-
tents are not always solid ; they frequently exhibit one
or more cavities; they are even frothy and granular,
like the contents of the nuclear utricles and the cells.
To these we may add the analogy with animal nucleoli,
which are likewise utricles, and the other utricular struc-
tures of the cell-contents.

* Nageli on Riiizocarpese, Ipc. cit.

f Nageli on the Moving Spiral Pikments (spermatic filaments) in Ferns.
(Bewegliche Spiralfaden, &c.) Schleiden u. Nageli's Zeitschrift f. wiss.
Bot., Heft i, p. 174 (1844).

t Nageli on Cells, &c. ; Ray Translation, 1845, p. 250.

UTRICULAR STRUCTURES. 173

The nucleoli occur inside the nuclear utricles and the
spermatic utricles.* Nothing is known of their mode
of origin. In animal nucleoli a process of division has
been observed, f

The shape of the nucleoli is globular ; at the moment
when they originate by division, hemispherical. If they
lie upon the membrane of the nuclear utricle, they are
sometimes flattened on that side. The contents of the
nucleoli are originally homogeneous and mucilaginous,
and usually exhibit greater density than those of the
nuclear or spermatic utricles ; in rare cases they are less
dense, causing the nucleolus to appear like a hollow
space. The homogeneous contents of the nucleolus sub-
sequently become frothy and granular ; or they remain
homogeneous and exhibit one or more cavities.

4. Mucilage- (Protoplasm-} utricles (Schleimblaschen).J

These structures have hitherto been either overlooked,
or taken for large mucilaginous granules or for globules of
agglomerated mucilage. They may be seen beautifully,
and in great abundance, in the Characese, where, together
with amorphous masses of mucilage, they are the chief
bodies seen to rotate. I have also seen them in the cells
of many Algse, of the leaves of Mosses and Hepaticee, in
the Perns and the Phanerogamia. They do not present
themselves constantly, and mostly but sparingly, 1 6
in a cell. They are more certain to be found in large
cells than in small.

The mucilage-utricles are distinguished from the mu-
cilaginous granules by their much more considerable size,
by possessing a perfectly spherical shape and a smooth sur-
face, and by refracting light less. That they are utricles,

* Nageli on Cells, &c. ; Ray Translation, 1845, pi. vii, fig. 11 a. Nageli
on Moving Filaments, &c., loc. cit. pi. iv, fig. 16, a, b, c.

j- KdlliSer, Entwick. der Cephalopoden, pi. vi, fig. 68.

5 I refer the reader to the observation made in the preceding essay
(p. 124) on the term " mucilage" (schleim).

174 . UTRICULAR STRUCTURES.

and not homogeneous granules, follows from this : 1st,
that when large enough, a membrane may be detected
on them which is not coloured by iodine, while the con-
tents become brown (pi. II, fig. 18) ; and 2d, that the
mucilaginous contents may become frothy and hollow.
The membrane becomes especially manifest, when the
contracting contents separate in places from the mem-
brane, in utricles which come in contact with water or
tincture of iodine (fig. 18, b, c).

Little can be observed of the origin of the mucilage-
utricles. They first appear as very minute globules of
mucilage. It is possible that a small quantity of muci-
lage becomes agglomerated, individualized, and acquires
a membranous covering. I have not observed a propaga-
tion of the mucilage-utricles.

The shape of the mucilage-utricles is spherical from
the beginning, and it always remains so. The growth
serves merely to expand the utricles uniformly.

The contents are originally homogeneous. They remain
in this condition, or one or more hollow spaces appear in
them.

The membrane is delicate, and seldom attains any con-
siderable thickness. In Chara and Nitella only have I
yet seen the mucilage-utricles with tougher membranes.
The membrane is also thin here originally, and either in-
visible or only indistinct. By the application of iodine
we may mostly succeed in seeing it more clearly. It then
appears distinctly bordered by two lines (Fig. 18, a). At
a later period minute points (V) appear on the outer
surface, which with the increase of size show themselves
to be little spines (c, e). In old utricles the spines dis-
appear, and the outer surface of the membrane is irregular
and uneven (d). During this the membrane has been
constantly increasing in thickness.

Looking for an analogy for the spines just described,
one might think of the cilia of the germ-cells of Vauclieria,
and notice that the mucilage-utricles of the Characere
likewise move (rotate), while the mucilage-utricles of other

UTRICULAR STRUCTURES. 175

plants have not this property. But the objection here
applies that the mucilage-utricles rotate already before
the spinous coat is observable on them. I do not think,
therefore, that this has anything to do with the motion,
and would rather conjecture that it is to be explained
in the same way as the outer membrane of the pollen-
granule.

5. Proliferous utricles (Brutblaschen.)

I give this name to those utricles in which, as in
Caulerpa, the chlorophyll- and starch-granules originate.*
I formerly called them " mucilage-cellules" (schleim-
zelkhen], but this name now requires alteration in respect
to the rest of the nomenclature. Metteniusf saw the
proliferous utricles in the radical hairs, and in the hairs
of the upper surface of the leaf in Salvinia.

According to Hartig } the chlorophyll-granules originate
in the so-called euchrome-cells, one kind of the so-called
ptychodal utricles. The manner in which the researches
were instituted, however, does not appear to me to afford
the requisite guarantee, for results which can be depended
on. It therefore still remains uncertain whether and
where the proliferous utricles are found in the Phanero-
gamia.

The proliferous utricles first appear as homogeneous
globules of mucilage. When they become larger, we
detect a membrane on them, and mucilaginous contents,
coloured yellow or brown by iodine. According to
Mettenius, the utricles originate as minute amorphous
granules. If this be intended to signify those minute
homogeneous globules of mucilage, and not actual gra-
nules, it agrees with my observations. The globules
differ from the mucilaginous granules by their regular,
perfectly spherical form, their wholly smooth surface, and

* Nageli on Caulerpa, Zeitschr. fiir wiss. Botanik., Heft i, pp. 184 190.
f Beitrage zur Kenntniss der Uhizocarpeen, p. 51, pi. ii, figs. 42, 43.
I Das Leben der Pflanzcnzelle, pp. 8-10.

176 UTRICULAR STRUCTURES.

the smaller power of refracting light. The exact process
of the origin of the proliferous utricles is as yet unknown.
Perhaps minute portions of the homogeneous mucilage of
the cell become individualized, acquire a globular shape,
and produce a membrane on their surface. Propagation
has not yet been observed in the proliferous utricles.

The proliferous utricles lie free in the contents of their
cell, and are always of globular shape. They increase in
size up to a certain point.

The contents of the proliferous utricles at first consist
of homogeneous mucilage ; this frequently becomes gra-
nular. In it originate several globules of starch- or chlo-
rophyll-granules.* The membrane of the proliferous
utricles is subsequently dissolved, and the starch-globules
and chlorophyll granules lie free in the cavity of the cell.

6. Colour-utricles (Farbblaschen).

These include the chlorophyll-granules, and the other
coloured globules of the cell-sap. They have already
here and there been called utricles, but no membrane has
been shown to exist upon them. Formerly, the starch-
globule, mostly present in their interior, was frequently
taken for a cavity.

The following facts prove that the said structures are
actually utricles: a whitish membrane, surrounding the
green contents, may be clearly seen in the larger chloro-
phyll-utricles of the Algae, of the leaves of Mosses, of the
pro-embryo of Ferns (pi. II, fig. 10 a, b, c), of Characese
(fig. 17), and, in favorable cases, in the leaves of Phane-
rogamia (fig. 12 a /). When they lie close together,
they do not become blended into a mass, but, like cells,
acquire a parenchymatous form (fig. 11). By abnormal
alteration (in the pro-embryo of Ferns (fig. 10 d, ,/),
in the leaves of Mosses and Hepaticse), they become

* On Caulerpa prolifera, by C. Nageli ; Zeitsclirift. f. wiss. Bot., Heft i,
1844, pi. iii, fig. 19.

UTRICULAR STRUCTURES. 177

larger; the contents lose their colour, and pass into a
colourless fluid containing granules ; in this condition they
are undistinguishable from minute cells. The contents
of the chlorophyll-utricles undergo peculiar changes, which
chiefly consist in the origin of one or more starch-globules
(fig. 12). Green and red colour- vesicles manifest growth,
and in the course of this their shape is altered in various
ways. Finally, the chlorophyll-utricles divide like cells
(% 10 c).

Since Mohl's researches* the chlorophyll- utricles have
commonly been regarded as a nucleus of one or more starch-
granules, on which chlorophyll has been deposited as a
coating. This view presupposes that the starch-granules
exist first, and that they are afterwards coated with chloro-
phyll. But the transition of starch-globules into chloro-
phyll-granules has nowhere been seen. On the other hand,
chlorophyll-utricles not unfrequently occur in the Algae
without a trace of starch in their interior. In some Algae,
moreover, as well as in higher plants, the origin and growth
of the starch-globules may be traced, in the interior of the
chlorophyll-utricles, up to the absorption of these latter.

The colour-utricles are best named according to their
colour ; green, red, blue, or yellow colour-utricles, or even
with one word ; green utricles (chlorophyll-utricles), red
utricles, &c.

I know nothing of the origin of colour-utricles free in
the cell-contents. They appear as minute (green or red)
granules, which, after they have reached a sufficient size,
display an utricular structure (fig. 10, a, b; fig. 12; fig. 17).
They originate by the division of a parent-utricle. This
becomes elongated, divides by a wall, and separates into
two new colour-utricles (fig. lOc).

I have seen the division of the green colour-utricles in
Algae (e. g. in Bryopsis Balbisiana, Valonia ovalis], in
the pro-embryo of Ferns, and in Nitella. This division
sometimes presents such an appearance that we only see

* Untersuchung. iib. die Anatom. Verhaltnisse des Chlorophylls. (Dis-
sertation, 1837, Verm. Schrift. p. 349.)

178 UTRICULAR STRUCTURES,

a circular constriction, which advances inwards, and at
last parts the chlorophyll-utricle into two. In other cases
a septum is first perceived, and the apparent constriction
is recognised to be simply a result of the retraction of the
already-formed secondary utricles from each other. Thus
a conviction is readily attained that in the first case the
septum is overlooked on account of its delicacy, or its
oblique position.

It may, indeed, be objected that the said division is
merely apparent, and produced artificially by the close
apposition of two colour-utricles. Such a condition is
actually produced artificially, when two utricles come so
close together under the microscope, that they become
flattened by their mutual pressure. It is therefore a
question whether to take account of that apparent division
of the chlorophyll-utricle in the vegetable-cell, or not.
Nitella has furnished me with evidence for it.

I examined the terminal cell of the leaves in various
stages of growth in the same individual, in Nitella syn-
carpa, namely, 1st, one measuring '080 of a line ; 2d, one
= '500; 3d, one =1*5; and 4th, one = 6 lines in length.
The diameters were from '030 to -050, '060, and '090
of a line. The chlorophyll-utricles lay in vertical rows
on the walls. They were all of about equal size, and ex-
hibited a perfectly regular shape, both in the young and
in the old cells. Some appeared to be in the commence-
ment of the act of division. Actual division and propa-
gation must take place if the number of chlorophyll-
utricles increase from the young to the old cells, since of
any very minute utricles which might originate free
among the others, I saw no trace, either between the
utricles of the same row, or between the rows. Now I
counted the rows, and found constantly about eighty of
them, both in the young and old cells. No multiplication
of the rows takes place ; in fact I did not see any chlo-
rophyll-utricle divide perpendicularly into two utricles lying
side by side. On the contrary, I found in the first and
shortest of the four cells mentioned, 40 utricles in a row ;

UTRICULAR STRUCTURES. 179

in the second cell, 150; in the third, 500; and in the
fourth, and longest cell, 2000. During the growth of
the cell from a length = *080 of a line to 6 lines, the
number of chlorophyll-utricles had increased in each row
from 40 to 2000, and in the whole cell from 3200 to
160,000. Consequently, while the cell became seventy-
five times longer, the chorophyll-utricles increased* about
fiftyfold.

I found the like in the cells of the stem of the same
plant. In a length of 1*3, and a diameter of '14 of a
line, I counted about 160 rows, in each row about 325
utricles ; in one about 20 lines long and *2 in diameter,
again about 160 rows, but some 3500 utricles in each
row ; and lastly, in a cell 30 lines long, and of a diameter
= '24, I found about 160 rows, and some 6700 chlo-
rophyll-utricles in each row. Here also the utricles
were of tolerably equal size, and in regular arrangement
in the cell. Individuals exhibited transverse division ;
but I saw no young, minute utricles between the others,
so that here also the increase in number could only be
effected by division. This increase amounted to about
twenty times the number, while the cell increased about
twenty-three times in length.

If the colour-utricles originate free in the cell-contents,
their shape is at first globular ; if they are produced by
the division of a parent-utricle, at first hemispherical or
semi-ellipsoidal. In the latter case they soon assume a
globular form as they separate from each other. If the
utricles, as rarely happens, remain free, so that they swim
in the cell-contents, they retain their globular form.
Usually they apply themselves upon the wall, whereby
their form is more or less altered, since they become more
or less flattened. Their section is then either almost
round, oval, hemispherical, serai-elliptical, or flatly com-
pressed.

The alteration of form which the colour-utricles undergo
during their growth consists not only of a mere flattening
of greater or less extent, which we see in the side view

180 UTRICULAR STRUCTURES.

(or section), but also in the acquirement of a different
shape, as seen in front. Here also the utricles at first
appear round. They either remain round or become
elliptical. If they lie close together, they become paren-
chymatous (fig. 11, fig. 13). If colour-utricles increase
in length, this takes place either parallel to the long axis
of the 'cell, or if they lie in a circulation thread, in the
direction of the current. I have observed the latter in
the Algae (e. g. in Conferva glomerata marina). On the
inner surface of the cell-wall lies a network of muci-
laginous filaments. The chlorophyll-utricles form simple
rows in the lines of the reticulation (fig. 16). They extend
in length in the direction of the lines, and may become
lanceolate, or even linear. Those lying in the angles of
the network assume a triangular form; the angles are
more or less drawn out in the direction of the filaments
of circulation towards which they are turned. In Floridese
(e. g. in Ceramium diaphanum) I saw the red utricles, which
were originally round, elongate so much in the direction
of the longitudinal diameter of the cell, that they had the
appearance of long fibres. In this case their transverse
diameter diminished considerably.

The growth of the colour-utricles presents great simi-
larity to the universal growth of cells. It exhibits on the
one side the transition from the globular to the tabular
form, on the other from the globular to the elongated,
fusiform or filamentous shape. They may also become
elongated into separate processes, almost like stellate cells.
Moreover, by pressure they may become parenchymatous.

The expansion which is connected with the growth of
the colour-utricles exhibits great quantitative differences.
It is greatest in those which originate free in the cell-con-
tents, since they appear first as very minute granules. It
is less in those which have been produced by the division
of a parent-utricle ; here it amounts, as a rule, to two to
three times the original length and breadth ; otherwise the
growth is not always connected with an increase of size
in all diameters ; in tabular and very much flattened

UTRICULAR STRUCTURES. 181

chlorophyll-utricles, the smallest diameter is shorter than
it was at first ; in the thin, fibre -like red utricles both the
shorter diameters are very considerably diminished.

The contents of the colour-utricles are at first always
homogeneous, green (fig. 12 a; 13 a\ red, yellow, or
blue. They may present different characters during the
life of the utricle.

In the first place, the contents remain for the whole
period homogeneous and coloured, without alteration, or
at least without any that can be noticed.

Or, secondly, one or more very minute granules become
visible in the contents, which are permanent in this stage,
and the nature of which cannot be made out, on account
of the small size. Probably they are starch- globules, as
would result from the following. In a third condition of
the contents, minute granules of the same kind appear,
but are somewhat larger, and appear as whitish globules,
coloured violet or blue by iodine (fig, 10, a, b, c ; fig. 11 ;
fig. 13; fig. 17). Mohl thinks that the violet tinge is
produced by the brown colouring of the surrounding
green contents. It appears certain to me, however, that
this is proper to the starch-globules. Even before the
treatment with iodine a difference is observable in the
latter, for they refract light more strongly and the others
more weakly. If they refract light in this manner, they
will acquire the blue colour, in spite of the surrounding
chlorophyll ; if the refraction is less, the colour is violet.
These starch-globules are very frequently solitary in an
utricle ; sometimes, however, from two to five are present.
They only occupy a small portion of the cavity of the
utricle.

A third condition of the contents consists in the fol-
lowing : As before, minute granules first become visible
in the homogeneous, coloured contents. They increase
in size, and at last almost wholly fill the utricles, so that
merely a thin layer of colour remains investing the starch-
globule (fig. 12 i). If, as is generally the case here, the
membrane of the utricle is indistinct, it appears as though

182 UTRICULAR STRUCTURES.

the starch-globule were only coated with colouring matter
(chlorophyll) .

A fifth condition. One or more starch-globules ori-
ginate in the colouring matter (fig. 12, af), which in-
crease in size, and finally almost fill the utricle (fig. 12,
ff /). The colouring matter (chlorophyll) by degrees
vanishes, at last entirely, and finally the membrane also
of the utricle is dissolved, and the starch-globules lie free
in the cell (fig. 12, m, n}.

These are the principal alterations which take place in
the contents of the colour- utricles. I have chiefly ob-
served them in chlorophyll -utricles. Nothing general
can be said respecting the distribution of the forms men-
tioned. As a rule, all or many are met with in the same
Natural Order. I have, however, met with green utricles
with contents remaining homogeneous, and those minute
starch-globules, coloured blue by iodine, much more fre-
quently in the Algae and Mosses ; those with large starch-
globules, coloured blue by iodine, more frequently in the
higher plants. The red-utricles of the Florideae gene-
rally exhibit only homogeneous red contents. The yellow
and blue utricles I am not yet sufficiently acquainted
with.

It still remains to be noticed that globules occur, in
rare instances, in the place of the starch, inside the colour-
utricles, which are either not coloured at all by iodine, or
become yellow. In my opinion they are corn-posed of
a substance analogous to starch ; allied to gelatine and
inuline. But I cannot agree with Schleiden, who says,
that chlorophyll (the chlorophyll-globules) often invests
starch, and also often other substances.* Beyond starch
(as ordinarily), and the globules just mentioned (which
occur exceptionally), I have never yet met with any
known solid product in the chlorophyll-utricles, for in-
stance, not even oil- or mucilage-granules, &c.

The homogeneous colouring matter of the colour-utri-

* Grundzuge, 2d ed. i, p. 190.

UTRICULAR STRUCTURES. 183

cles also undergoes an alteration occasionally, consisting
in the colour becoming fainter; moreover, even in the
transition of one colour into another. In many genera
of FlorideaB it is rather common for the red colour-utri-
cles to become subsequently green or yellowish green.

The colour-utricles undergo dissolution in three ways :
1, by propagation ; 2, by the starch-globules inclosed
within them gradually displacing the colouring matter,
and at length causing the absorption of the membrane ;
and, 3, by changes in the cell-contents, inducing a solu-
tion of the contents and membrane.

When the solution of the colour-utricles is brought about
by changes in the cell-contents, they usually become de-
tached from the cell-wall, pass into the interior of the cavity
of the cell, and acquire a more or less perfectly globular
form. The colouring matter becomes granular, loses its
colour, and is then dissolved, the membrane at the same
time disappearing. In decaying cells of the pro-embryo
of the Ferns, as well as occasionally in decaying cells of
the leaves of the Hepaticse and Mosses, I have seen the
chlorophyll-utricles expand considerably during this pro-
cess (in Pteris nemoralis from "006 to *008 of a line).
The chlorophyll dissolved into minute dark granules which
swam in a colourless fluid. The membrane of the utricle
was here very distinct. (Fig. 10, d, e,f.)

7. Starch-utricles, Starch-granules.

Three different opinions have been expressed as to the
origin and nature of the starch-granules. According to
one, they consist of a cell-like envelope and fluid con-
tents. According to a second, a nucleus is first formed,
around which concentric layers are deposited on the out-
side. According to a third view, the formation of the
layers proceeds from without inwards.

The first opinion, Raspail's, appears in a peculiar form,
resulting from his theory of cells. The starch-grain is a

I 'S I I TKK'ULAK STIUTCTUKKS.

cell or an utricle originally attached to the wall of the
((ill, and produced by this. The envelope is starch; the
contents gummy (gurmnos) and seiniilnid. Microscopic
investigation is in opposition to this view.

The second opinion, as champions of which I'Yitsche
and Schleiden may be named, maintain, on chemical
grounds, the fundamental point, that, with the exception
of UK; nucleus, the layers are wholly composed of the
same chemical substance. The outermost layers are im-
pregnated with foreign matters, and arc thus rendered
insoluble in water. Especial dillicnlties occur in this
view, both in chemical and physiological respects, in ex-
plaining the nucleus, which, according to the authors
mentioned, on one hand, is a hollow space, on the other,
contains neither sugar, gum, nor starch. What then is
the thing upon which the first and innermost layer of
starch is deposited.

The third opinion has been recently promulgated by
Munter.* The innermost layers are the softest, and
thus the youngest. Very good ; but the origin of
the outermost and earliest layers of starch is here
forgotten ; upon what are these deposited ? Of course
there is no difficulty about those succeeding. More-
over, a fact which Munter states that he has ascer-
tained in the starch-grains of Gloriosa superba, stands in
strange contradiction to this view, namely, " that an or-
ganic compound also may appear in a crystalline form ;"
and then the term of " starch-concretions" (starkedmsen)
proposed for these so-called agglomerated granules is just
as inapplicable.

In my opinion, the starch-grains are utricles, and con-
sist, like the other utricles, of a membrane arid lluid
contents. Concentric layers are deposited on the inside
of the membrane as in lignifying cells. The cavity of
the utricle (the so-called nucleus) thus becomes reduced
to a most minute excavation, which is always filled with
fluid.

* Holanisfhe Zcitung-, 1815, j>. I '.:;.

UTRICULAR STRUCTURES. 185

The grounds for the opinion here expressed are as fol-
lows : In many starch-grains a layer different from the
rest, and surrounding the nucleus, may be more or less
clearly perceived. In some, this shows itself as a tolerably
thick membrane which iodine does not colour (fig. 14).
The starch-grains are hollow ; sometimes the cavity is so
large that the starch invests the membrane merely as a
thin layer (fig. 15). The layers become softer and con-
tain more water toward the interior, and are tougher and
more solid toward the exterior ; from analogy to the lig-
nifying layers of cells, the softer are to be considered as
the younger, the harder the older.

Nothing is known of the origin and propagation of
the starch-utricles, except what regards the external condi-
tions under which they are produced ; we know that they
may be formed free in the cell, and this either in mucilage
or in chlorophyll ; in utricles, namely, in the interior of
nuclear utricles (in the Fucoideas*), of prolific utricles
(Caulerpd), and of colour-utricles (pi. II, fig. 10, 11, 12,
13, 17).

The shape in which the starch-utricles appear is pro-
bably at first always spherical. They may remain spheri-
cal or extend lengthways, so as to become almost
cylindrical, or grow in a plane, so as to assume a tabular
shape. The form of the full-grown starch-grains is ex-
ceedingly varied and mostly irregular, so, however, that
granules which originate free, are always bounded by one
single curved surface.

The granules lie free or united in groups of from two
to twelve, and even more. A group of this kind is, not
very aptly, termed a compound grain. The individual
grains or utricles are parenchymatous ; the sides directed
towards the interior of the group are plane, those which
form the surface of it are curved (convex outwards). All
the grains united together in a single heap have been
produced in one utricle. They are at first spherical, but
from the reciprocal pressure which they undergo, in con-
sequence of their growth, they acquire a parenchymatous

* Nageli on Cell-Formation ; Part I, llay Trans. 1845, pi. vi, fig. Ifi.

186 UTRICULAR STRUCTURES.

form, and through this same pressure adhere together as
a tissue (fig. 12 g). If the utricle then becomes absorbed,
the grains either remain united in a group (fig. 12 n) or
separate. Since, however, they have acquired a solid
structure by the formation of layers of lignification, they
do not reassume the globular form, but remain polygonal
and exhibit a shape which has a crystalline appearance,
without, on that account, being actually crystals (fig, 14).

I do not mean, however, that the starch-grains which
have originated together in the same utricle, necessarily
always assume a parenchymatous shape ; they only do
this when they are contained in proportionate number or
size (to the size of the utricle). If they lie more loosely
in the utricle, they always remain isolated and retain their
rounded surface. Thus I have never seen starch- grains
become properly parenchymatous, either in the nuclear
or prolific utricles. In the green utricles, also, I have
likewise often seen them remain globular, even when a
number have originated together in one utricle.

Those starch-grains, therefore, which lie separate and
are bounded by a single curved surface, have originated
either free in the cell, isolated in utricles, or in numbers
loosely arranged in utricles. Those which lie separate
and have one or more plane surfaces or angles, have
originated in numbers, closely packed in a green utricle.

In like manner the granules united into a mass have
been formed together in a green utricle.

The original contents of the starch-utricles are unknown.
As soon as they have attained a sufficient size to allow of
their being investigated, they are coloured blue, violet, or
red, by iodine. The formation of layers, therefore, begins
very early, and it must thence be assumed that the outer-
most and first layers expand considerably in surface until
the growth of the utricle is completed. There is nothing
whatever opposed to this hypothesis, since the layers are
at first in a soft, semifluid condition, and since the cell-
membranes and the earliest of its layers of thickening are
likewise capable of considerable expansion.

Consequently starch -utricles and cells agree in this: that

UTRICULAR STRUCTURES,

187

the membrane of both begins to be thickened soon after
its production. They are distinguished by the fact that in
the starch-utricles this thickening is relatively much more
considerable, even at the very beginning, than in the cell.

The structure of the starch-utricles first becomes clear
when they have acquired a greater size ; this is : a mem-
brane, probably composed of gelatine, layers formed of
starch, and a cavity filled with watery fluid. The mem-
brane is more or less distinct ; when it is thick enough
we may perceive that it remains uncoloured in the treat-
ment with iodine, while the layers become blue or violet
(%. 14 15).

The layers are formed like the thickening layers of the
cell ; the concentric lines are to be explained in the same
way as these. The layers are composed of starch through-
out their entire thickness. They decrease in solidity
from without inward, and contain more water in the
reversed order. Hot water and acids cause them to swell
up very much, and they frequently tear in consequence
of an unequal expansion, either merely in the interior, or
from within outward. But that an air-bubble originates
in the cavity through the action of sulphuric acid, as has
been asserted, I have not seen, neither can I conceive how
sulphuric acid could produce an air-bubble here, by ab-
straction of water. For the acid does not draw out the
water like some pump or exhausting apparatus, bat when
the acid is applied to the starch-grain a reciprocation of
currents is set up, according to the law of endosmosis
and exosmosis, through the layers and the membrane, the
acid passing in, and a portion of the fluid of the cavity
and the layers going out. If more fluid passed out than
in, and the starch-utricle could not contract in a corre-
sponding degree, the only consequence would be that the
diminution of density, beginning in the interior, would
cause an increased flow inwards from without. Now the
swelling up of the utricle proves exactly the contrary,
namely, that more fluid passes in than out.

The cavity of the starch-utricle is sometimes larger,
sometimes smaller. In a given size of the utricle of

188 UTRICULAR STRUCTURES.

course its size is in inverse proportion to the thickness of
the layers. In many grains the cavity is reduced to a
minimum. These grains correspond to the cells of the
stony concretions of pears, and completely lignified liber-
cells. In other grains the cavity is of considerable size.
The thickness of the deposited layer of starch may be so
small that the starch-utricle resembles a cell with mode-
rately-thickened walls (fig. 15). I found such a condition
not unfrequently in the pith and rind of the fruit panicle
of Vitis. The cavity is filled with an almost colourless
fluid, which doubtless is no simple chemical substance,
but contains formative matter, such as gum, sugar, and
quaternary compounds, although in small quantity.

8. General Retrospect.

We have found various organic structures in the cell-
contents possessing, in all their phenomena, a great
resemblance to the cell itself. The general term for those
structures, partly on account of this similarity, partly to
express the actual difference, may most fitly be utricle.

The utricle agrees with the cell* in the following
characters. It probably originates by the isolation of a
(minute] portion of organic substance, which becomes coated
with a membrane. Therefore from the very origin there
appears a distinction between contents and membrane.
The utricle grows both in its membrane and its contents,
and, in the course of this, changes its shape in manifold
ways. The membrane expands and becomes thickened by
the deposition of layers in the interior. The contents are
metamorphosed, and produce new organic forms. Finally,
the utricle propagates.

Thus nothing occurs to the cell which is not found in
the utricle, at least in one or a portion of the kinds of
utricle. Moreover, we see that exactly the essential
peculiarities of the cell, the proper membrane and the
transformable contents, occur in all utricles. The identity
between cell and utricle is, therefore, tolerably manifest.

* See the preceding Essay (on Cell-Formation).

UTRICULAR STRUCTURES. 189

The distinction between the two is more difficult to
find, especially because the origin of the utricle is not yet
sufficiently made out. I believe, however, that even with
our present knowledge, the distinction may be at least so
far established, that cell and utricle appear as absolute,
distinct conceptions. The comparison may take into
consideration either the relation of the two structures to
the individual plant, or the two structures independently
of general relations.

In reference to the relation to the vegetable organism,
cell and utricle exhibit the following distinction. The
cell is the elementary organ, which takes part immediately
in the formation of a tissue. The plant first becomes
independent with the cell. It developes through the
formation and growth of cells ; it lives through the cells.
The utricle, on the other hand, is an elementary organ
which takes part only mediately in the origin, structure,
and life of the plant, because the utricle is merely a part
of the cell. Consequently we may call the cell the imme-
diate, the utricle the mediate elementary organ of the plant.

Considering cell and utricle independently, as indivi-
dual organisms, we can at present establish merely the
following distinction. The cell needs for its production
a nucleus, formed previously ; it is formed around a
portion of organic cell-contents in which a nucleus is
inclosed.* The utricles, on the contrary, never originate
around a nucleus, only the nuclear utricles originate
around a nucleolus. We may accordingly define the
cell and the utricle in the following terms :

The cell is an individual quantity of contents, inclosed
by a homogeneous membrane, and individualized through
the influence of a nuclear utricle ; it is the immediate
elementary organ of the vegetable organism.

The utricle is an individual quantity of contents, in-

* I have shown, in the preceding Essay, that in the first place a nucleus
is produced which individualizes a greater or smaller quantity of the sur-
rounding contents, through attraction, and that this individualized portion
of contents acquires a membranous coat upon its surface.

190 UTRICULAR STRUCTURES.

closed by a homogeneous membrane, and individualized
without the influence of a nuclear utricle ; it is merely the
mediate elementary organ of the vegetable organism.
The following kinds of utricle may be distinguished :

1. Nuclear utricle, Nucleus. It originates under the
influence of a nucleolus, and contains, at least in the earlier
periods of its life, scarcely anything, excepting one or more
nucleoli (colourless or coloured), besides mucilage. It
occurs (with the exception of those which originate in a
parent-cell) only immediately in the cell (not in another
utricle).

2. Spermatic utricle. It originates, probably, like the
nuclear utricle, in like manner, under the influence of a
nucleolus. It contains principally mucilage, and produces
a spermatic filament in its interior. The spermatic utricle
probably occurs originally merely in cells, and in any case
never inside other utricles.

3. Nucleolus. It contains mucilage, and occurs (some-
times with the exception of those around which the
nuclear utricle has not yet been formed) only inside the
nuclear utricle.

4. Mucilage-utricle. It contains mucilage, but no
nucleolus, and occurs immediately in the cell (not in other
utricles).

5. Proliferous utricle. It at first contains mucilage,
but no nucleolus, subsequently colour- or starch-utricles.,
which, after its solution, become free. It occurs only free
in the cell (not in other utricles).

6. Colour -utricle. It contains colouring matter, but no
nucleolus, and may be named according to the colour.
It frequently contains also starch-utricles. It occurs free
in the cell as well as in old nuclear and proliferous utricles.

7. Starch-utricle, starch-granule. It contains layers of
thickening of deposited starch, and occurs free in the cells,
as well as in old nuclear, colour, and proliferous utricles.

ANNUAL REPORT

ON RESEARCHES IN

PHYSIOLOGICAL BOTANY,

DURING THE YEARS 1844 AND 1845.

BY DR. H. F. LINK,

DIRECTOR OP THE KOYAL BOTANIC GARDEN OP BERLIN,

PHYSIOLOGICAL BOTANY.

GENERAL OBSERVATIONS.

PHYSIOLOGY, and with it Physiological Botany, were, it
was thought, quietly making considerable progress, since
the number of contributors continually increased, and
though of course opinions differed, none were maintained
with any remarkable violence. Then certain individuals
appeared who endeavoured to disturb this tranquillity,
and not only advocated their own opinions with great
violence, but attacked those who thought differently,
challenged them, and even in some cases insulted them.
I shall especially mention three of these Liebig, Gaudi-
chaud, and Schleiden. All three write well, Liebig
excellently so ; not one is wanting in genius and inge-
nuity; in their zeal, however, they have not all been
able to command themselves, but have given way to a
violence which, although not hindering them for any
time, perhaps even aiding them in acquiring speedy re-
nown, is nevertheless always injurious to the cause which
they are anxious to advocate.

Liebig says, in the first edition of his celebrated book,
' Organic Chemistry in its Application to Agriculture and
Physiology' (Brunswick, 1840), p. 35, " As soon as phy-
siologists meet with the mysterious vital force in any
phenomenon, they renounce their senses and faculties ;
the eye, the understanding, the judgment, and the re-
flective faculties, all are paralysed as soon as a phe-
nomenon is declared to be incomprehensible." Now
this has not really been the case ; they have indeed very

194 PHYSIOLOGICAL BOTANY.

rarely declared a phenomenon to be incomprehensible,
whilst they have frequently erred in the contrary direc-
tion ; but supposing they had done so, they could always
say confidently to those philosophers who wish to believe
that everything depends upon mechanics, and forces
which act mechanically " Tell us, ye slanderers, are you
then acquainted with the fundamental theory of the whole
of your mechanics; do you understand the cause of
motion even in the slightest degree ? Is it not the most
incomprehensible of all the phenomena with which we
are surrounded?" And even if it were answered that it
was the first, the most common, and the most certain
empirical knowledge upon which anything could be based
with certainty, it might be easily replied, that the same
was also the case with vitality, for we cannot even start
the question of the cause of motion without living. What
has just been stated might be recommended to many
philosophers, especially in foreign countries, as a subject
for careful consideration, when they urge mechanical ex-
planations to the very utmost when, as it were, they
are suspended in the air without any support. Dutrochet
may serve as an example ; he attempts to explain me-
chanically all the motions of plants, by means of endos-
mose and exosmose, by the influx and egress of the fluids
in the cells and vessels which permeate the membranes,
fill and distend the cells, and produce movements by
means of this distension, and even by flowing out pro-
duce collapse and motion in the opposite direction. Yet
the causes of the phenomena of endosmose and exosmose,
upon which this theory is based, have by no means been
ascertained; it has certainly not been proved that the
exchange of matters in solution through the unvitalised
membrane, which we find in these experiments, occurs
through the living membrane of the cells in plants, for
the simple reason, that we do not find that those cells
which are situated near each other contain different fluids,
by means of which this exchange could be effected ; we
cannot understand how the gradual influx and egress

PHYSIOLOGICAL BOTANY. 195

occurring in endosmosis and exosmosis can produce
rapid movements, especially those of Mimosa pudica, to
which Dutrochet applies his theory ; lastly, it cannot be
explained by mechanical laws how this expansion and
contraction of the cells becomes capable of raising whole
portions of plants. Still this is repeated by the majority,
and De Candolle was once at their head ; but I must
cease here, to avoid falling into the very error that I have
attributed to others.

Is it not better, instead of retarding the progress of
science by such explanations and their details, to recur at
once to a vital force, the determination of the laws of which
will remain our object, and our not unfounded hope ?

To a certain extent, but to a certain extent only, has
Liebig returned to the ordinary path in regard to the
vital force. In his book, * Organic Chemistry in its Ap-
plication to Physiology and Pathology' (translated by
Dr. Gregory, London, 1842), in the third part, speaking
of the phenomena of motion in the animal organism
(p. 196), he says : " If the vital phenomena be considered
as manifestations of a peculiar force, then the effects of
this force must be regulated by certain laws, which laws
may be investigated ; and these laws must be in harmony
with the universal laws of resistance and motion, which
preserve in their courses the worlds of our own and other
systems, and which also determine changes of form and
structure in material bodies, altogether independently of
the matter in which vital activity appears to reside, or of
the form in which vitality is manifested." The author is
by no means clear in his views of the pretended vital
force. What is meant by " must be in harmony ?" Are
they the same, or do they only resemble each other ? We
do not see why they might not be directly opposed, or
totally different. But the author's views are not clear
even upon a purely physical force, the attraction of
gravitation. He says (1. c. p. 201) : " If it (the stone)
fall from a certain height, it makes a permanent impres-
sion on the spot on which it falls ; if it fall from a still

196 PHYSIOLOGICAL BOTANY.

greater height (during a longer time), it perforates the
table ; its own motion is communicated to a certain num-
ber of the particles of the wood, which now fall along
with the stone itself. The stone, while at rest, possessed
none of these properties. The velocity of the falling body
is always the effect of the moving force, and is, c&teris
paribus, proportional to the attraction of gravitation. A
body falling freely acquires, at the end of one second, a
velocity of 30 feet. The same body, if falling on the
moon, would acquire, in one second, only a velocity of
sloths of a foot, = 0*1 inch, because in the moon, the
intensity of gravitation (the pressure acting on the body,
the moving power) is 3600 times less."

We shall not dwell upon the individual expressions
which are not always accurately applied, but shall merely
ask, Why does Liebig omit to notice the law of inertia,
upon which all mechanical determinations are based,
which causes the velocity of a falling body to increase
constantly in proportion to the duration of its fall. Galileo
applied it without recognising it ; when he discovered
the law, that the space through which a body falls is as
the square of the time in which it falls. Newton called
it the law of inertia, placed it at the head of his ' Principia
Philosophise Naturalis Mathematica/ and expressed it as
follows : " A body continues in its state of rest or motion,
in the same direction and with the same velocity, until
some motive force compels it to change this state." It
may be perhaps attributed to the physio-philosophers
of Germany that they have forgotten, or at least over-
looked, this law in the explanation of natural phenomena ;
in fact, that natural philosophers, who, like Liebig, have
no respect for philosophy, make no mention of this law.
Not only does it explain the increased velocity acquired
by a falling body, but even the most common, the daily
phenomena occurring upon the moving globe, could not
be explained without it such, e. g. as the fall of a stone
from a house or a tower; why, when allowed to fall
from the west side of a house it does not reach the earth

PHYSIOLOGICAL BOTANY. 197

far behind the house ; because the house being situated
upon the globe, which is in rapid motion, flies away as
it were from it ; and lastly why, when a stone falls from
a high tower, a deviation ensues, because the summit of
the tower is in more rapid motion than the earth at its
foot, towards which the stone descends. But I feel
ashamed to be teaching matters which belong to ele-
mentary school-instruction. Newton correctly ascribed
to matter inertia, and not a property of passive resistance,
as some philosophers attribute to it ; for so long as the
matter is subjected to this law, it has no property ; it is
in a perfect state of indifference; it cannot assume a
state of motion when at rest ; it cannot, in the slightest
degree, alter any motion which has been imparted to it
from without, and without its co-operation ; in short, if
we may use the expression, it is lifeless. Here, then, we
have a certain and definite character of inactivity, from
which we may start, and from which we must start, in the
consideration of vitality and the vital force. The anti-
thesis of vitality, in contrast with this state of inertia,
this indifference, is clear ; a body must be called living
when capable of spontaneously assuming a state of motion
from one of rest, or when capable of changing or in any
way determining its motion ; whence what is to be con-
sidered as the vital force soon follows. Let us take one
application of what we have stated : Is the universal at-
tractive force a vital force ? The answer is negative ; the
body approximates to another, merely inasmuch as it is
attracted ; it does not then set itself in motion ; it does not
determine its motion by any power of its own ; this is
merely determined by the attraction of some other body.
Hence alone is astronomy enabled to calculate with cer-
tainty and accuracy the motions of the celestial bodies.
This force can certainly set other bodies in motion, but not
that body in which it exists, and through which it acts.
We cannot in any way understand why there should not
be forces capable of setting the bodies in which they exist
in motion, since we find motions in living bodies which

198 PHYSIOLOGICAL BOTANY.

cannot be derived from external forces. These are de-
nominated vital forces. They are not in the least more
incomprehensible than the force of attraction ; in fact
they are less so. Newton found, with regard to the force
of attraction, that the intensity with which it acts upon a
body is inversely proportional to the square of the distance
of the body from the centre of attraction. But has this
law been shown to exist in the case of other forces ? Does
it apply to cohesion, elasticity, magnetism, &c. ?

Liebig says, in his ' Chemical Letters ' (Heidelberg,
1844, p. 18), "Medical writers and kindred minds are
annoyed at the great simplicity of the truth, although
they have not succeeded in practically applying it, not-
withstanding all their pains ; they therefore give us the
most improbable views, and by the term vital force they
understand a wonderful thing, by means of which they
explain everything which they do not understand ; by a
thoroughly incomprehensible and indefinite something,
everything which is incomprehensible is explained ! "

That bodies combine chemically in definite proportions
only is undoubtedly a law of combination, but I should
like to know according to what law chemical decom-
positions take place. Is the chemistry of decomposition
anything more than a specification of the results of ex-
periments made upon individual bodies, each individual
experiment being calculated according to a fixed form ?
Is the word affinity anything more than a word ? Nothing
is explained in chemistry; everything in chemistry is
incomprehensible. The vital force, on the other hand,
acknowledges general laws. There is the law of pe-
riodicity, which is the direct antithesis of the law of
inertia, according to which motion increases to a certain
extent, and then again diminishes ; there is the law of
habit, according to which reaction is not always equal to
action, but diminishes in proportion to the frequency of
the repetition of reaction. However, I am not writing
upon general physiology.

Liebig's work, mentioned above (Organic Chemistry

PHYSIOLOGICAL BOTANY. 199

in its Application to Physiology and Pathology) is still
an excellent one. It shows us how the proportion of the
constituents of the fluids, and of the solids of the animal
body, may be deduced from the proportions of the con-
stituents of the nutritive matters, This is certainly the
first step towards explaining the nutrition of the animal
body and its secretions ; but it is only the first step, and
we are ignorant of the decomposing and the combining
forces. They appear to belong to physical rather than
vital forces ; and even when we have ascertained these,
the ultimate means by which these forces are set in action
remain to be determined. And for the physician, the
principal point is to increase the activity of these forces ;
or when it is too great, to diminish it. We must agree
with Liebig, nay, we might even censure with his violence,
when we see how many physiologists misapply the word
life ; but this blame does not attach to all physiologists,
when they used the term vital force correctly, i. e. when
they apply it to those cases in which chemical forces cease
to act in their natural manner. It is very necessary to
advance by means of physics and chemistry as far as
possible, but we must not trust more to these two
sciences than they are capable of effecting.

Gaudichaud has come forward with great decision
against Mirbel in the Academy of Sciences at Paris.
Offended by some expressions which Mirbel made use of
in his ' Memoir upon the Structure of the Stem of the
Date-palm/ and which Gaudichaud correctly considered
as directed against him, immediately after the reading of
the memoir, he briefly protested against it, and declared
Mirbel' s system to be incorrect. There also appeared
soon after, in the year 1843, two memoirs justifying his
protest. They have been spoken of in the 'Annual
Report ' for 1842-3, as have also his EechercJies generates
sur T Organographie, &c. des Plantes in the ^Annual
Report ' for 1841. In the ' Comptes Rendus ' for 1844,
we now have the third and fourth protests against Mirbel
(I, pp. 597 and 899). He has not yielded. In 1844

200 PHYSIOLOGICAL BOTANY.

Mirbel read a memoir upon the Structure of the Stem of
Dracaena Australis, of which we shall speak hereafter ;
and in the ' Comptes Rendus ' for 1845, we find not less
than seven papers against Mirbel's memoir. Many years
ago even, Gaudichaud was so engrossed with his own
theory, that he would scarcely listen to objections made
orally to him, or which referred to investigations which
he was going subsequently to publish. His style is brief,
almost aphoristic, and positive, nevertheless it is not free
from repetition ; but he says, by way of exculpation,
" I must continue to repeat until my system is generally
received." Prom his obstinacy it may be expected that
he will not yield.

Gaudichaud has also evinced this pertinacity in his life.
He was an apothecary, accompanied, as pharmaceutical
botanist, a voyage of discovery under the command of
Freycinet, and, in September 1817, went on board the
corvette Urania, which was shipwrecked on the 14th of
February, 1820, at the Maluines. It sailed from Port
Jackson southwards, next struck upon some ice-islands,
then rounded Cape Horn, and cast anchor in the Bay of
Good Success, at Terra del Fuego. A violent gale of
wind compelled them to cut their cable and stand out to
sea. Some days afterwards, during the most beautiful
weather, the ship was driven upon some concealed rocks
near the Maluines, and twelve hours afterwards upon the
sand in the bay or creek called by the French the Bay
of Solitude, where it still remains. It was four o'clock
in the afternoon when it went upon the rocks, and four
o'clock in the morning when it was wrecked on the sand-
bank and sunk. The interval between these two events
was a dreadful night of anxiety and danger. Gaudichaud
was fortunately saved, but the whole of his collections
were submerged, and could not be removed until they
had been from thirty-six to forty hours under water. He
was obliged to wash every packet, in fact every sheet, in
fresh water, and to dry them ; and thus during the four
months which he passed there, he was enabled to save

PHYSIOLOGICAL BOTANY. 201

about 4000 specimens of plants out of 6000. He re-
turned to France in December 1820, with the corvette
La Physidenne, which the government had purchased at
the Maluines. He wrote there the botanical part of the
description of the voyage, and also planned his 'Organo-
graphy and Physiology of Plants.' In 1831 he went in
the frigate Nerminie, under command of Villeneuve
Bargemont, to the coast of South America. The frigate
sailed twice round Cape Horn, and returned in 1832
from Rio de Janeiro to France. Gaudichaud, however,
obtained permission to remain in the Brazils, from which
country he returned to Toulon in the corvette La Bonite,
Captain Durand, in June 1843. In April 1835, he de-
livered his remarks on the f Organography, Organogeny,
and Physiology of Plants ' to the Institute ; and in Decem-
ber of that year, on the day on which the Monthyon-prize
was awarded to him, he left Paris to make his third voyage
in the corvette La Bonite. He started from Toulon in
February 1836, and returned in the same ship at the
end of the year 1837. I have derived this information
from the account of his life in the ' Revue generale Bio-
graphique,' which adds further : " Gaudichaud, that
energetic man, born with the Revolution in 1789, and
grown up in it, has fought several duels, but," it adds,
" all who were acquainted with M. Gaudichaud know
that he never took the lead in these affairs." I have not
given these details of Gaudichaud' s life here entirely
without an object.

"What," says he (Compt. rend., 1844, 1,598) "is a
Monocotyledonous plant at its very origin, e. g. a Date-
palm? an animated cell, which produces an embryo or a
bud. An embryo, as all botanists now know, is a free,
isolated, independent cell. This embryo, or this primi-
tive phyton, is a distinct individual, possessing its own
peculiar organization, and its peculiar functions. The
first individual soon produces a second; the second, a
third ; the third, a fourth, and so on, during the whole
life of the plant. As the embryo possesses its organiza-

202 PHYSIOLOGICAL BOTANY.

tion and its peculiar normal functions, so will also the
individuals produced by it, and by all those which follow
it, and which possess their own separately, i. e. modified
according to the stages of their development and their
age, the second being directly and constantly grafted upon
the first ; the third upon the second, and so on, each one
being grafted upon the other. The first individual, the
embryo, derives the principles of its existence from without,
from water, air, light and heat, but especially from the
albuminous body (perisperm), when it is present, which
nourishes the embryo, and is thus absorbed ; the second
is nourished by the first ; the third by the second and
the first, and the fourth by the three others, as well as by
the elements previously named; hence it foUows, that
when the phytons are perfectly developed, the first re-
mains very weak ; the second is somewhat stronger ; the
third is still stronger ; and that all the subsequent ones
become stronger and stronger, as also more complicated
in form, and consequently in functions, until we arrive at
the normal leaf, which has attained the highest stage of
organization."

Gaudichaud says, all botanists now know that the em-
bryo is a bud, whence it may be readily deduced, that
the bud is just the same as the embryo, and must have
roots like it. But this is not the case, the embryo is not
a bud, nor the bud an embryo. There is an old and
every-day observation, which I am accustomed briefly to
explain in the following manner : the embryo produced
by impregnation propagates the species ; the bud, the
individual. A branch with buds, from a Borsdorf apple-
tree, when grafted always produces Borsdorf apples,
whilst the seed of a Borsdorf apple never does so. Dif-
fering in this important property, it may also differ in
other respects, hence it does not follow that the bud
should have roots like the embryo. Moreover, the leaf is
not an individual, it is only such when united with the
buds, and these at first contain only cellular tissue,
extremely few spiral vessels. Such buds, when united
together, form Mirbel's Phyllopliore.

PHYSIOLOGICAL BOTANY. 203

We shall allow Gaudichaud to continue.

" According to the old theories, the vascular system of
the second individual is formed by a division of the
vessels of the first. Hence the vascular system of the
second individual is composed of a portion of that of the
first. But if the vascular organization of the second in-
dividual is more compound than that of the first, the
vascular system of the second cannot be formed from
the vascular system of the first. If it be granted that all
the vessels of the embryo pass into the primordial leaf,
the latter should always possess the organization of the
embryo only. But this theory is, I think, now properly
rejected. According to the theory propounded to you
on the 12th of July (by Mirbel), the vessels of the primor-
dial leaf must emanate from the inner periphery of the
embryo. We are here met by the same difficulties. In
fact, what becomes of this theory, when it is proved to
you by a large number of facts that, as a rule, the primor-
dial leaf is more highly organized than the embryonal
leaf, and that e. g. the fourth and fifth leaf almost always
contain more vessels than the three or four first leaves ;
moreover, when we show by the same facts, not only that
the cotyledonary leaf does not send vessels to the pri-
mordial leaf, but also, that in many cases it does not
receive any from above, and it then has only an ephemeral
existence. In this case the first leaf ceases to exist, be-
cause it is not strengthened, and to a certain extent
vitalised by the second leaf. Is not this an evident proof
of the individual vitality of the phyton?"

Mirbel maintains, if I am correct, that all the vessels
of a Palm-stem are not only derived from the internal
periphery of the embryo, but also, that wherever leaves
arise, new vessels are developed, and thus that they appear
at the rings, internally, at the periphery of the stems.
From observations which I have made upon this point, I
believe that Mirbel is incorrect ; 1 do not find bundles of
woody or vascular tissue arising from the rings within
the stem, but they all appear to arise from the base of

204 PHYSIOLOGICAL BOTANY.

the stem, and then traverse it in its entire length. Near
the periphery the vascular bundles are so crowded that it
is difficult to separate them and ascertain their course.

Gaudichaud continues : " We shall naturally apply this
principle to the growth of the stem, leaves, fruit, &c., and
also extend it to the flowers and other deciduous parts of
plants. We would also apply it to the stem of Vellosia,
which, as it derives scarcely anything from the leaves
which exist at the ends of the branches, always remains
very thin, for the simple reason that the vessels of the roots
of the leaves which should produce the increased thickness
of the stem, become applied immediately after their for-
mation to the external part of the bark (a Vexterieur du
perixyle), and thus descend as roots (a Tetat de ratines),
along the twigs, the branches, and leaves, to the ground.
The primordial leaf (that first formed after the embryo),
undoubtedly obtains life and nutriment, but nothing fur-
ther, from the embryo; the primordial leaf also imparts
vitality and its principal nutriment to the second leaf, and
the same occurs with the second leaf in regard to the
third, &c."

Gaudichaud always writes in this aphoristic style,
which becomes more remarkable from his separating the
sentences from each other, and commencing a new line.

Again, he says (1. c. p. 610): " In fact, since it is found
by observation, that the embryo, this small isolated being,
consists originally of cellular tissue only, and that this
cellular tissue produces the vessels by its physiological
action ; that the vessels commence in the internodes of
the stern (merithale tigillaire), then appear in the petiolary
and the laminary merithals of the leaf; that they are per-
fectly formed, or at least may be traced in these regions
(dans les parties merithattiennes), before they appear in the
papilla of the rootlet (mammelon radiculaire), we are led
by analogy to consider that the same must also be the case
with the organization of other individuals, of whatever
kind they may be, which are produced by plants. This
fact I repeat is an important one, and worthy of conside-

PHYSIOLOGICAL BOTANY. 205

ration. I have several times recurred to it, and shall
again recur to it, because I believe that it is the key to
vegetable organography ; because it comprises a theory of
the regions (or members) (merithalles), which I defend;
and because it supersedes (infirme) all other theories."
The basis of his system is principally contained in these
words.

Gaudichaud's whole theory rests upon the perfect
identity of the buds and the embryo, and the formation
of roots or root-like parts in the former as in the latter,
when the branch grows. This explains the increase in
thickness, which is always a difficulty, especially in the
Monocotyledons, and in the cauloma of Palms. It has
already been stated above, that the resemblance of the
embryo and the buds is not so great as Gaudichaud thinks.
Roots growing downwards in the stem are attributed to
buds from analogy only, and this is imperfect. Although
Mirbel explains the increase of the stem of Palms by the
origin of new vessels from the interior of the circum-
ference of the stem, yet the accurate explanation of the
entire process is accompanied with great difficulties, in-
dependently of the fact that this origin of the vessels
cannot be found on accurate examination. But the
difficulties vanish if we admit the occurrence of lateral
growth, which appears probable at first sight. In my
Lectures on Botany (Part II, Berlin, 1845, p. 309), I have
shown that the stem of the Date-palm in its early stage,
much resembles a bulb, which also at first increases in
thickness, and then ascends in the form of a stem ; I have,
moreover there (p. 237) detailed observations, which show
that in the stem of the Dicotyledons, a layer, sometimes
thick, sometimes thin, applies itself; this, however,
especially indicating lateral growth. Thus the roots of
the buds do not pass far down the stem. It is extremely
probable that a little cellular tissue may grow downwards
from the buds into the stem, but it is doubtful whether
vessels descend from the buds into the stem, they certainly
do not pass down far in this manner (see my Lectures, &c.,

206 PHYSIOLOGICAL BOTANY.

p. 265.) Mirbel and Gaudichaud pay altogether too little
attention to the relative growth and position of the vessels
as regards each other. The pertinacious man, as evidenced
in his life, will hardly sacrifice any of his theory to con-
troversy; but if it should ultimately tire even the impartial
critic, this ought never to go so far as to cause the theory
to be rejected without examination.

Schleiden is the most violent of the botanists mentioned
at the commencement. As soon as he meets with an
opposite opinion, he rejects it immediately, and so posi-
tively, as not to admit the correctness of a single assertion
in it. The author of such opinions fares still worse;
everything of his ceases to be good. By this means, with
few exceptions, he has excited all botanical writers against
him, and caused many of his own theories to be rejected
by others. This should not prevent the acknowledgment
of what is good and excellent in him. When we see the
straightforward and positive Gaudichaud, we expect a
pertinacious adherence to his opinions, but from Liebig's
amiable aspect we should not anticipate so severe a man,
nor should we suspect, of the quiet Schleiden, that he
could trample upon all who differ from him. The first
edition of Schleiden's ' Principles of Scientific Botany' was
not incorrectly called a libel, in France ; this reproach
would be unjustly made to the second, on the whole,
although in parts the same violence is manifest, and pro-
bably expresses his feeling. After its dedication to
Humboldt, which all who know Humboldt must admire,
the following passage immediately follows, in the preface :
" It is an infinitely difficult task, again, to throw off the
accessory means of education and to retain education
alone, to apply the stifled power independently to objects
which have been spontaneously selected. On the large
scale this is most strikingly seen in the laughable pre-
judice for Latino-philological erudition, and the mediaeval
monkish book-wisdom, which makes all true living pro-
gress in our education appear distorted and crippled as
an inherited dyscrasy, and just where it appears in its

PHYSIOLOGICAL BOTANY. 207

greatest absurdity, in the Natural sciences, still disturbs
the fresh springs of life." Had this been said one hun-
dred years or more ago, it might have been regarded as
an expression made at an appropriate time, but it now
comes much too late. We must now rather thank those
men who, like Humboldt, appreciate the advantage of
still cultivating the taste for the dead languages and
philological erudition. Humboldt has done this in very
many of his writings, very recently in his ' Cosmos/ in
a manner which, as we may hope, and must wish, will
exert its influence upon an age which is too fond of what
is superficial and easy, beyond which it has no desire to
proceed. I shall allude here to the effect upon the mind
which the wonderful force and simplicity of the dead
languages exert, when we regard the impression which
they produce only, without referring to the dilution
which they always suffer by translation into modern
languages. This is foreign to my present purpose. But
in the Natural sciences their use is really not absurd, it
is much to be recommended in descriptive Natural
History, and hitherto has always been retained.

In these languages all European nations understand
each other ; the plants and animals described by us,
Germans, are again recognised from Lisbon to Moscow.
Schleiden speaks of the absurdity of species-trifling ; but
here again, in his peculiar style, he says too much, be-
cause the first point is to ascertain what is spoken of,
and the determination of species must serve as the alpha-
bet of science > and it then leads us to the answer to one
of the most important questions in Botany, viz. what is a
species, what a variety, and how the latter are produced ?
It is, perhaps, most proper that papers which dispense
with this, as it were, mechanical method of illustrating
objects, should be wholly written in the mother tongue ;
but it would be well if so much Latin were learned as to
cause those manuals which make use of an aphoristic
style of description, especially in the Natural sciences, to
be understood in every foreign country. The English,

208 PHYSIOLOGICAL BOTANY.

French, and Italians, even at present, are but little ac-
quainted with our labours in the Natural sciences. We,
with whom it is part of the education of youth to learn
the languages of these nations, more readily become ac-
quainted with the labours of other nations than they do
with ours, because our language is much too difficult to
be learned by these nations. Hitherto the Russians, in
their writings on these sciences, have generally made use
of Latin, French, and German ; but if they were to begin
to write in their own languages, and simultaneously to
make great progress in the sciences, we must then either
remain in ignorance of their works or learn their lan-
guage. But Schleiden condemns learning from books,
and in his opinion it is of no consequence whether we
are acquainted with what has been observed by foreigners
or not. He says, in the same preface, " True formative
knowledge, that which expands the nobler powers of the
human mind, can at the most be acquired in and with
the assistance of books, but never from books. Learning
from books is the secret and unsuspected source from
which the disingenuousness and tendency to dissimula-
tion is first nourished, and which poisons the present age :
accustoms us from youth upwards neither to say, to
think, or to do anything of ourselves, but merely to fill
our meagre and sterile minds with ideas which have been
borrowed and handed down from other sources, so as to
palm off this abundance as healthy knowledge/' He
frequently repeats that he has striven to be peculiar and
original. In the same preface he says, " I had endea-
voured at once, and without any regard to what had
already been done, but furnished with all the expedients
at our command in the present day, to rediscover the
whole science from the direct investigation of nature;
thus my work acquired an originality, which, independ-
ently of its correctness, always possesses something more
attractive than the matter when arranged in an historical
and philological form." The author somewhat deceives
himself. Where there is a noise, there flock boys and

PHYSIOLOGICAL BOTANY. 209

idlers. He is far less original in his views than Thouars,
Turpin, Agardh, Nees von Essenbeck, or Oken, and in
description, Gaudichaud is far more vigorous and distinct.
As regards his accuracy, this cannot be so easily and
readily decided upon, as to exert any striking influence
upon the judgment of the reader. Thus, e. g. in the
first edition of his book, he follows the theory of the
French botanists on the stem, which he certainly defines
with more minuteness ; and in the treating of the stem of
the Palm, he criticises what I have said upon the subject,
but without supporting his remarks by any original ob-
servations. Original writers are certainly not those who
have produced most benefit to science, whilst on the other
hand they have frequently retarded its progress, and I
should not consider it as any recommendation, were any
one to assert that Schleiden was original in his botanical
remarks. On the whole, he recommends the critical
method, in fact, considers it as the only correct one ; but
we cannot possibly conceive criticism without some
preceding system ; in fact, it is quite opposed to pecu-
liarity and originality. It is highly valuable, and we
should be grateful to the author's ingenuity if he allowed
his criticisms to be decided and severe, but free from such
extravagancies, which impair rather than increase the
effect they produce. The observations of any writer upon
natural history, who only describes what he has seen, are
very valuable ; but it would be impossible to re-institute
a science from the study of nature alone. In making
investigations we must know what to observe, and this
must be acquired from instruction, and finally from
books. Without these means we should merely make
discoveries which had long been known. Had it not
been learned from books, we should not have been aware
that iodine colours starch blue. I was .deprived of this
resource in my earlier researches, and it has since proved
of much service in science. It is too great an exaggera-
tion, nay, it is even false, to assert that books cherish a
disingenuousness and tendency to dissimulation which

210 PHYSIOLOGICAL BOTANY.

poison our whole existence. We might with more truth
accuse social life altogether, which no doubt frequently
renders dissimulation necessary to prevent one being
thrown upon the street (obliged to beg).

The author, whilst combating dogmatism in his ' Me-
thodological Elements' (p. 23), passes the following incor-
rect judgment upon Endlicher and linger' s ' Principles of
Botany' (Vienna, 1843). "This false plan is carried out
to its utmost extent in the recent work of Endlicher and
Unger, and its appearance under the protection of siich
names can only be seriously regretted. It appears to me
that independently of many of the details being objec-
tionable, to which we shall allude hereafter, the authors
in writing then- book in a rigidly scholastic style, at the
present day, have committed a great mistake. From
beginning to end, it contains mere explanations of names
arranged systematically, and what renders them especially
useless, is that the authors have rarely taken the trouble
to name examples. Anatomy, physiology, and the history
of development, which alone should constitute the peculiar
value and true foundation of the details, are very meagre
and unimportant, the figures, which are appended at the
end, are neither formally nor essentially brought into
connexion with the details which are deducible from them
only." All knowledge in the natural sciences depends
upon definitions, for every fact is comprehended as a
definite conception. Merely because the perception of an
object or occurrence is repeated, does it become absorbed
as manifold, in the unity of the idea, and in this form, we
become acquainted with it. In all sciences, and espe-
cially in natural history, we must commence with defini-
tions. We must first obtain a definite idea of a part of
an organic body ; the external form and the connexion
with other parts are the first and the most important
points to be regarded, for by them we recognise the part ;
the internal structure, the anatomy, certainly must be
known, but it is entirely a subordinate matter. Then
follows the doctrine of development, for I must first know

PHYSIOLOGICAL BOTANY. 211

what is developed, and from what this is derived ; and the
physiological examination of the species follows last. Now
I confess that I am unacquainted with any manual of
botany which, with aphoristic brevity, fulfils its design so
admirably, as the 'Principles of Botany' of Endlicher and
linger. My disagreement with the authors in a few, in
fact, in many of their theories, is nothing to the point, for,
in so extensive a field, it is impossible always to meet
with accuracy. Schleiden finds fault, e. g. with the dis-
tinction which the authors make between a conical and a
discoid receptacle, when speaking of the floral receptacle,
and puts a variety of questions, which in my opinion may be
easily answered. The discoid receptacle is furnished be-
neath the ovary with an annular ridge, which is absent in
the conical receptacle, and if I understand the authors
correctly, they regard this as an indication of another
inter node of the stem which commences there. Thus they
have explained the presence of the various parts situated
beneath the ovary, for by explanation we signify the illus-
tration of the essential connexion of phenomena. But I
doubt whether an appendage does not always exist beneath
the ovary, indicating the origin of another internode.

Schleiden' s theory of the internodes of the stem, meri-
thalles, as they are called by the French in their usual
manner, by a barbarous term derived from the Greek and
in opposition to all analogy, is old. The place where a
leaf and a bud exist, was called a node, and this was
regarded as the commencement of an internode. In the
Grasses, each node is evidently the commencement of an
internode ; in the Palms, internodes are closely crowded
and somewhat less easily recognised ; the nodes and the
internodes are also distinguishable in the Labiatse, the
Caryophyllacese, &c., which have opposite leaves, whilst
in plants with alternate leaves, they run into each other.
If we consider the term node to denote the articulation,
we might say with Endlicher and linger, that in the
conical receptacle, there is no node situated above the
stamens, until we come to the ovary, whilst in the discoid
receptacle, one does exist.

212 PHYSIOLOGICAL BOTANY.

" The peculiarity of the inductive and modern method,"
says Schleiden (p. 25), "consists in our first completely
abstracting ourselves from all hypothesis, not premising
any principle, but starting immediately from our direct
consciousness, from individual facts, endeavouring purely
and completely to isolate these, arranging them according
to their essential affinity, ascertaining from them alone
laws to which they are subject, and which are essential to
their existence, and thus tracing them back, until we
arrive at those ultimate ideas and laws, with which all
further deduction ceases to be possible." This may be
very true ; but it is least applicable when the doctrine of
development, anatomy, and physiology are assumed as
the foundation of the investigation. The second book,
the doctrine of cells in plants, begins as follows (p. 197) :
" Cells can only be formed in a liquid which contains
sugar, dextrine, and mucilage (cytoblastema). It takes
place in two ways : 1st. The mucilaginous parts become
condensed into a more or less roundishbody, the cell-nucleus
(cytoblast) and at their whole surface convert part of the
liquid into gelatine, or comparatively insoluble substance; a
closed gelatinous vesicle is formed, the external liquid
penetrates into and distends this, so that the mucilaginous
body becomes unattached on one side, and remains ad-
herent to the other side of the internal walls; it then
forms a new layer on its free side, and is thus inclosed in
a duplicature of the wall, or it remains free, and is then
usually dissolved and disappears. During the gradual
distension of the vesicle, the gelatine of the wall usually
becomes converted into cellular matter, and the formation
of the cell (cellula) is completed. 2d. The entire con-
tents of the cell become divided into two or more parts,
and from each a delicate* gelatinous membrane is imme-
diately formed ; in this manner several cells are simul-
taneously developed, which then completely fill the cell in
which they were formed."

Upon how much that is uncertain is this based ! In
his explanation, the author himself directly says, " We
are far from clear as to the liquid from which the cells

PHYSIOLOGICAL BOTANY. 213

are formed." This is so certain, that the author com-
mences with " It appears." It is moreover uncertain,
and is doubted by many, myself among the number,
whether a cytoblast is formed before the surrounding
membrane ; we have never seen it. Although we may
find granules, and subsequently cells, in a clear liquid, it
does not follow that the former are formed from the latter,
moreover, the young cells under these circumstances are
frequently empty ; sometimes certainly they contain several
nuclei. Moreover, it is hypothetical and cannot be seen,
that the nucleus of the cell converts part of the liquid
into gelatine ; that the external liquid permeates the gela-
tinous vesicle and distends it, is also hypothetical -, and,
lastly, it is no less hypothetical that the gelatine of the wall
becomes converted into cellular matter, and that in this
manner the cells are perfected. It is by no means my
intention to assert, that these facts are false; I merely wish
to say, that we must not commence with these statements
with what is doubtful and uncertain.

I have several times reminded the reader that the cells
of the Alga3 cannot be regarded as analogues by which
the development of cells in the Phanerogainia can be
explained. The cells of the Algae are rather to be com-
pared to the joints of the stem in the Phanerogamia, than
with the individual cells of which the stem is composed.
The cells of the Algae are placed in a long tube, and
hence were called utriculi, and, in fact, utriculi matricales
by Roth. Moreover, the remarkable phenomena which
are perceptible in several of the cells of the Algae, as,
e. g. in Spirogyra, Stellulina, &c., appear to characterise
them as peculiar organs. The author remarks, in passing
(p. 205) : " To guard against false views I must observe
here, that the theory of crystallization brought forward
by Link, according to which crystals are formed by the
confluence of small globules, depends upon imperfect
observation." I never thought of saying anything of the
kind. When a recent precipitate, as e. g. of carbonate of
lime, is quickly placed under the microscope, we observe

214 PHYSIOLOGICAL BOTANY.

nothing but globules, and what proves them to be in a
fluid state, is that they are frequently seen to run into
one another. The crystal is then suddenly formed ; in
the above instance, it is a rhombohedron or crystal of
Arragonite, according to the temperature. Schleiden has
not seen this, and is unacquainted with my little work, * On
the Formation of Solid Bodies/ Berlin, 1841. My friends
H. and G. Rose and Poggendorf have seen it. But
further : "At first it is natural, that if we desire to
observe the formation of crystals, precipitation should
not be selected for the purpose, which chemists consider
to belong to the so-called irregular form of crystallization,
but that the observations should be made at first upon
simple crystals whilst separating from concentrated liquids.
In this way we observe in each case, as e. g. with nitres,
ammonio-chloride of platinum, and most beautifully and
readily with the ammonio-chloride of zinc, &c., that the
nuclear crystal suddenly springs up, not at any given
moment, in the liquid which was previously perfectly
clear, and which remains so, and then whilst apparently
at perfect rest, is seen gradually to increase in almost
imperceptible starts by external deposition." If some
chemists consider precipitation as only irregular crystalli-
zation, they have done wrong. The above means of
making the observations are totally impracticable. If the
concentrated solution be allowed to evaporate slowly, the
formation of the crystals can only be observed with great
difficulty ; if it be allowed to cool suddenly, the crystals
are formed so suddenly and in such numbers, that indi-
vidual crystals are difficult to trace. Precipitates which
crystallize slowly are the best to examine, as e. g. carbonate
of lime, very little of which must be placed under the mi-
croscope. In precipitates which crystallize rapidly, as sul-
phate of lime, the first stage of globules cannot always be
perceived, as the crystallization ensues too rapidly ; some-
times, however, for this very reason, the phenomenon is
seen with surprising distinctness." Further : " If, how-
ever, two liquids which yield a single precipitate be mixed

PHYSIOLOGICAL BOTANY. 215

under the microscope, at the instant of their coming into
contact, the sudden formation of a membrane, which
separates the two liquids, is seen to occur. On minute
examination, this membrane is seen to consist of crystals,
some of which can be perceived with perfect distinctness,
some are seen to be crystals when magnified more strongly,
others only when the very highest magnifying powers are
used, until lastly the smallest appear only as points, even
with the very highest powers. If the liquids are not dis-
turbed, some of the crystals which are formed gradually
enlarge on both sides in the liquid ; but if the liquids be
mixed, a large number of the crystals are instantly redis-
solved, others continue to increase in size, and fresh crys-
talline nuclei are suddenly formed in spots where the
liquid was perfectly clear." These observations are upon
the whole correct ; the so-called membrane is a wall of
turbid liquid. As long as it appears like a membrane, it
is not composed of crystals, but these are soon formed,
and it is then composed of them. A similar turbid wall
is also seen when the freezing of water is observed with
the microscope. See Poggendorff's Annal., vol. 64 (1845),
p. 479. Lastly, " After having made many observations
and these with great care, I have arrived at the general
conclusion, that every inorganic substance, when passing
undisturbed into the solid state, immediately assumes a
crystalline form ; most of the so-called pulverulent preci-
pitates consist of crystals, and the comparatively minute
size of others prevents our giving any opinion upon their
form." This is certainly the common opinion. But
Ehrenberg was the first to show that many fossils consist
of globules arranged in rows, and therefore are not com-
posed of crystals, and if the drop of liquid containing
the precipitate of carbonate of lime, under the microscope,
dries up too quickly, a quantity of powder remains be-
tween the rhombohedra, and this entirely consists of small
globules. The pulverulent condition of matter therefore,
which, I believe, Weiss is almost the only one to admit
as a distinct state, cannot be rejected. That crystals are

216 PHYSIOLOGICAL BOTANY.

not at once formed in the liquid, but that a nucleus is
first suddenly formed from a liquid, which subsequently
increases in size, my observations upon precipitates have
shown distinctly.

The author's remarks (p. 53 et seq.) upon the produc-
tion of the various forms occurring in nature are upon
the whole correct and to the purpose. The form either
excludes the mother-liquid, i. e. the formative fluid, during
its production, or it incloses it. The first is the case in
inorganic, and the latter in organic bodies. I should not
say, that the crystal during its formation excludes the
formative fluid, for the entire globule, or the entire aggre-
gation of globules, in the above experiments is transformed
into the crystal. Moreover, this consideration appears
to contradict the author's own opinion upon crystalliza-
tion, according to which the crystal is at once formed in
the liquid, and during its increase merely withdraws
particles from the formative liquid. But it is certainly of
great importance that organic bodies should be formed
within an envelope, where external agencies are directed
towards the centre of the formative liquid. When the
author says : " We thus characterise the idea, organism,
as the relation of the figure or form to the inclosed
mother-liquid, and life as the reciprocal action exerted
between the mother-liquid and the form," he must, on a
little reflection, understand how very unsatisfactory are
these characteristics. It was, therefore, with satisfaction
that I read the author's remarks (p. 64 et seq.) upon
Minerals, Plants, and Animals. They contain if he will
not take it unkindly a poetical effusion, which if it does
not distort the facts, forms an agreeable embellishment
to the subject.

The treatise on the Microscope (p. 82 et seq.) may be
strongly recommended to those who make use of this
instrument, although I find the following passage at the
end of it (p. 105) : "It is considered that little more is
required to make a microscopic observation than a good

PHYSIOLOGICAL BOTANY. 217

instrument and an object ; the eye has then only to be
kept above the eyepiece for the observer to be au fait.
Link, in the preface to his Anatomical Plates, expresses
this thoroughly false view as follows : ' I have generally
intrusted the observations entirely to my draughtsman,
M. Schmidt, and thus the unprejudiced condition of the
observer, who is unacquainted with botanical theories,
vouches for the accuracy of the drawings/ The result
of this perversion is, that Link's phytotomical plates, not-
withstanding his celebrated name, are so useless, that the
beginner, at least, must be strongly warned in learning
from them, to avoid his being confused by representations
which are entirely false. Link might as well have asked
a child, or a congenitally-blind person who had just been
operated upon, the apparent distance of the moon, and
from their freedom from prejudice, have expected the best
judgment, just as if in our early years of childhood we com-
mence learning to see with our unaided eyes," &c. I must
subjoin here the preface to my Anatomico-botanical Plates
(Pt. i, 1837) : " The anatomy of the human body only
first began to make the great advances in which it now
rejoices, when philosophers began to have the appearances
delineated by skilful artists. I followed this course as
far as I was able. For philosophers are seldom good
draughtsmen, and even when they do understand the art,
they have no time for its exercise. Hence it very fre-
quently happens that they draw what they have never
seen, or what they fancy they have seen under the mis-
guidance of some theory. This is especially the case
when the objects can only be seen under the microscope.
The most proper person for this purpose is a skilful artist
who is unacquainted with anatomical science, and who
must not be told what he is to see. A young artist,
C. H. Schmidt, who is a flower-pain ter, has for seven
years drawn for me the internal structures of plants, as
seen under the microscope. After he had become accus-
tomed to the microscope, I told him that he must only
draw what he saw, and always unhesitatingly contradict

218 PHYSIOLOGICAL BOTANY.

me, if he thought differently from me. He did not inte-
rest himself in theories, not even in mine. I have pre-
sented some figures selected from a large number, which
appear to me very accurately and carefully drawn, and
shall continue to do so if the undertaking should receive
support/' I therefore by no means leave the observation
to the draughtsman, but only the delineation ; I correct
him, but do not at once desire compliance, as with a
young and dim-sighted artist, but contradiction. I con-
fess that I had in my mind the Plates upon the Circulating
System of Plants, and especially Meyen's illustration of
the network of the so-called vital vessels in the leaves of
Alisma plantago. The brief preface to the second half of
the Anatomico-botanical Plates concludes with the follow-
ing words : " But we learn to see, both with the eyes
given to us by nature, and with those formed by art."
From that time to the present (January 1846), M. Schmidt
works with me five days every week in the morning,
except during my autumnal tour, and does not draw any-
thing which I have not carefully observed, and my eyes,
thank God ! are as good as ever. I educated my draughts-
man for microscopic delineation, and at the end of seven
years he was so far advanced, that I could reason with
him ; now after sixteen years he is still more so. How
can any one be thought so foolish as to have drawings
made under his own superintendence without pointing
out their object. I beg M. Schleiden not to consider
every one a fool but himself.

I must, however, apologize to the reader for having be-
come prolix on matters relating to myself. But some-
thing more upon a purely scientific subject. "That
property of the cell has already been mentioned," says
the author, in the chapter upon " The Life of .the Cell,"
p. 273, "in virtue of which it transmits fluids. It is
a perfectly superfluous and clumsy hypothesis, in the
explanation of this point, to have recourse to the exist-
ence of minute invisible pores ; the membrane and
liquid here stand in the same relation to each other, as

PHYSIOLOGICAL BOTANY. 219

salt and the water by which it is dissolved. Just as in
this case both salt and water are contained in every
differential of magnitude (sit venia verbo), so is the
cellular matter and the water in that of the membrane ;
with this difference, that the membrane is never rendered
fluid by the water, because the latter merely dissolves a
definite minute quantity, and then does not take up
more water until the portion first taken up has been
removed." Now, where do the aqueous particles exist
in the membrane ? They cannot exist anywhere but in
its interstices, however minute these may be, and how-
ever minute the particles of the membrane may be,
between which the aqueous particles penetrate. Un-
doubtedly such interstices, which we call invisible pores,
must exist, unless we admit an infinite penetration of
the water and the membrane. Independently of the
fact that such penetration cannot be perceived, nor even
imagined, the water and membrane would then form an
indivisible substance. The penetration would also be
mere groundless hypothesis. Salt, when in solution,
certainly exists only in the interstices of the water;
soluble bodies force carbonic acid from the pores of
water, because they take its place themselves. Our
system of physics must be altogether modified, if we are
to deny the existence of invisible pores. Physio-philo-
sophy is alone capable of affording an explanation of this,
because, according to its theorems, all forms of matter
are originally the same, and the differences depend upon
an increase or a diminution of the cohesive attraction
of one for the other. Still it would be difficult to find in
it an explanation, where membrane and water only are con-
cerned, without admitting the existence of such pores. Are
we, then, who are accustomed to work with the micro-
scope, to pretend that we can see everything? Thus
there are different gases, none of which are visible to us,
and in which we must admit the existence of large inter-
spaces, for the explanation of the phenomena which are
exhibited when they are mixed with each other or with
aqueous vapour. That these pores are not disseminated

220 PHYSIOLOGICAL BOTANY.

empty spaces, is readily understood ; but they are, in
most cases, filled with some subtile matters, as air, heat,
&c. The membrane of organic bodies allows fluids to
permeate it ; in endosmosis this is probably effected by
electricity. In the living body, these pores appear to
open and close an effect of the vital force, which in
many other cases manifests itself to us by contraction and
expansion.

Schleiden completely follows Fries in his philosophical
views, and has written a pamphlet against Hegel and
Schelling, in which he states that he does not attack their
system, but merely endeavours to show their ignorance of
natural science. The followers of these two philosophers
might find much to censure in it, as, in my opinion,
would the followers of Fries also, in his illustration and
application of the philosophy of Fries. I consider myself
as rather belonging to the latter ; but this is not the
place to discuss the matter. The author does not men-
tion Oken, who certainly deserves consideration. How-
ever, I will not give rise to a dispute, which under
the present circumstances could not prove of any advan-
tage to science.

Scientific disputes are usually beneficial to the pro-
gress of science. They not only augment the interest of
science itself by producing novelty in its monotonous
course, but they also possess the advantage of causing
the disputant to develope the grounds upon which his
opinion is based, more fully, for the sake of more clearly
illustrating it and convincing his opponent. Whether
he succeed in the latter or not, must be left undecided
by the disputant ; but so much is certain, that if this is
not immediately or very shortly the case, one or the other
ultimately acquires conviction. The advantage which
would result from the development of the grounds for or
against any view vanishes, when mere contradiction
i. e. rejection without a reason is made use of. It pro-
duces the least advantage to science when the dispute is
carried on with a truly original rudeness, as is generally
done by Schleiden.

PHYSIOLOGICAL BOTANY. 221

It is very wrong to repudiate physio-philosophy as
paying no regard to facts, but proceeding with purely
mental conceptions. Such is not the case. Oken, Nees
v. Esenbeck, and Wilbrand, like all other physio-philo-
sophers, assume facts as their basis; and only err, in
my opinion, in comprising them under ideas of too wide
an extent. Thus, under the idea of polarity, they
include so many heterogeneous phenomena, that the
definition and application of the idea becomes too arbi-
trary. In general, polarity signifies an antithesis in
different directions. This occurs very frequently in
nature, but so generally, that recourse to it not only
becomes tiresome, but even superfluous, and withdraws
us from more important and exact investigations. More
accurate and strict definition of the ideas is requisite, and
this again necessitates more minute and rigid determina-
tion of the facts. The opponents of physio-philosophy
have erred in this respect also. Thus the idea of a cell,
as now generally understood, cannot be mistaken ; but
when we find how the embryo-sac, the cells of the pith
and bark, spiral vessels and the joints of the Algae are
so comprised in this term, that what applies to one is
considered as holding good as regards the others, there
is risk of falling into the most serious errors. The
greatest injury effected by physio-philosophy, has arisen
from its not only rejecting mechanical philosophy, but
even holding it in contempt. Hence the fundamental
theories of physics, the theories of motion, have been
so neglected in courses of instruction, that we have had
to censure the want of acquaintance with them above,
even in the opponents of physio-philosophy themselves.

; INTERNAL STRUCTURE OF PLANTS.

In no department of physiological botany, if we except
the formation of the embryo, has so much been done
during the last few years, as on the formation and deve-

222 PHYSIOLOGICAL BOTANY.

lopment of cells. The investigations exhibit a depth of
research, which commences with the very earliest origin
of the plant, and in this respect they are very valuable.
First, Mohl, to whom we are indebted for most of our
knowledge on this point, has published :

Remarks on the Structure of the Vegetable Cell, in the
Botanische Zeitung. Edited by H. VON MOHL and L.
VON SCHLECHTENDAL. Berlin, 1844, p. 15 et seq., and
p. 273 et seq. He was led to these researches by
Hartig's investigations upon the structure of cells, and
his assumption of their possessing a more internal mem-
brane which lines their interior, and which he deno-
minated a Ptychode. "If we examine a first year's
shoot of a tree, or the stem of an annual plant, which,
before the completion of its growth, has been immersed
in spirit and kept in it for some time, we find in all
those cells and vessels, the secondary layers of which
have not yet attained their full development, an inner
membrane, which is remarkably distinct from the other
membranes of the cell. This membrane forms a per-
fectly closed, cell-like, thin-walled vesicle, which in the
fresh plant is closely applied to the inner wall of the cell,
and therefore escapes observation ; whilst in specimens
which have been preserved in spirit, it is contracted, and
more or less detached from the wall of the cell/' He
calls this cell-like vesicle the primordial utricle, and found
it in a number of Dicotyledonous plants, as, e. g, in
JSambucus ebulus, Ficus carica, Pinus sylvestris, Asclepias
Syriaca, Hoya carnosa, Euphorbia Canariensis, Caput
Medusa, &c. In the Monocotyledons, he detected it in
the apex of the stem and of the root. But this utricle
can be seen by a shorter method than that of keeping
the portions of the plants for a long time in spirit. In
general, the preparation need only be exposed to the
action of nitric or hydrochloric acid for a few minutes ; if
the acid be then neutralized with ammonia, and the pre-
paration be coloured by iodine, the primordial utricle is

PHYSIOLOGICAL BOTANY. 223

seen just as beautifully as by the long preservation in
spirit. As the primordial utricle exists in all young cells,
the author thinks that it contributes to the formation and
growth of the cells ; for, he adds, we can only conceive
cell-growth to occur in two ways, either by division of
the older cells by the formation of a septum, or the forma-
tion of cells within other cells. He believes that in the
cambium-layer of Pinus sylvestris, Sambucus ebulus, As-
depicts Syriaca, and Euphorbia Caput Medusa, he has
seen two primordial utricles prior to the appearance of a
septum between them, which, therefore, confirms the
latter method of formation. However, he is by no means
free from doubt on this point. What we have just
asserted entirely agrees with Schleiden' s theory, except
that Schleiden believes the cell-membrane to be formed
from the nucleus. Mohl, on the other hand, considers
that the cell-membrane always surrounds the nucleus.
Moreover, according to Schleiden, the cell-membrane first
formed constitutes also the later one the external mem-
brane of the cell; whilst, according to Mohl, the mem-
brane of the primordial utricle becomes the external
membrane. Hermann Karsten mentions having seen
the primordial utricle in his memoir, ' De celle Vitali ;' but
he confounded it with the internal layers of the membrane
of the cell. The author names several excellent exam-
ples of the various cell-membranes, and concludes, in
opposition to Hartig's opinion, as follows : " The above
remarks show that a positive decision of the question,
whether the cells are invested with a special membrane
or not, is accompanied with no little difficulty, since
optical illusion (Mohl means a luminous appearance) and
a slight modification in the substance of the innermost
layer of the cell, and this may also occur in the inter-
mediate layers, may readily lead to the belief that such a
membrane has been found. Hartig obtained his proof
from the cells of Taxus baccata, in which Mohl long
since showed that there existed a third layer. We must

224 PHYSIOLOGICAL BOTANY.

gratefully acknowledge that Mohl has now first taught
us the true structure of the membrane of the cell ; that
the wall of the cells and vessels is composed of a primary,
external, imperforate membrane, and a secondary one
which is usually perforated with apertures. It constitutes
the basis of our knowledge upon this subject. We may
add, with Payen, the outer membrane is not coloured
yellow by iodine, whilst the internal lining is so. Mohl
further adds, that the internal membrane consists of
superimposed layers. This is by no means rare, espe-
cially in the solid, cartilaginous, so-called stony cells,
several remarkable examples of which have been adduced
by the author also in this memoir ; but is not found in
all, at least has not been positively detected. Why, then,
should we admit their existence in parts where they are
not visible? How the primordial utricle is converted
into a cell having a separate existence, the author does
not by any means show; and we shall hereafter allude
to the fact, that it not only exists in the young cells, but
also in those which have completed their growth, and not
unfrequently even in old cells, provided they have not
become too solid and cartilaginous. But when Mohl
says, that the increase of cells takes place either by the
division of the older cells, by means of a newly -formed
septum, or by the formation of cells within other cells, a
third plan is evidently overlooked viz. the formation of
new cells between the old ones. Mirbel has already
shown this in his memoir upon Marchantia. It appears
to me to be the true manner in which their increase
takes place. I have had the anatomy of the bulb of
Amaryllis formosissima drawn in my Plates (Part I,
pi. 1). We there see, in fig. 4, at the base of the leaves,
a zone of short, laterally distended cells, with thinner
walls than those above and below them. Hence they
appear to have been recently formed ; moreover, the gra-
nules contained within them are not coloured blue by
iodine, as the granules in the cells, which are situated

PHYSIOLOGICAL BOTANY, 225

above and below them. The latter cells are large and
polygonal, tolerably uniform in diameter, and contain
large granules of starch. If we imagine to ourselves
these transverse cells extended longitudinally, they would
assume the same form as the polygonal cells existing
above them. These transverse cells appear to me to be
those last formed, and to have sprang up where the large
polygonal cells are separated from each other, and have
left gaps. That such gaps must be formed during the
growth of the parts is unavoidable. As the stem in-
creases in thickness, the woody bundles of the liber
become separated from the pith, and the layers of woody
tissue grow up between them. These certainly could
not force asunder the parts between which they grow,
but these latter must separate from each other, by virtue
of a distinct and peculiar vital power of expansion, to
allow the growth to take place. Physiologists, in giving
their attention to matters of little consequence, often
overlook others which are of great importance, as has
happened in the case of this peculiar property. Mohl
has nowhere shown, as he very modestly confesses, that
the increase of the cells is produced by the primordial
utricles.

In other general respects, the observations upon this
utricle, detailed by Mohl, are, as might be expected,
minute and accurate. I have not only examined plants
which have been long kept in spirits, but also, and much
more frequently, such as have been macerated for some
time in nitric acid. It is unnecessary to neutralize the
nitric acid with carbonate of ammonia, and the section
only requires to be washed with water, to obtain the
result just as distinctly. Colouring the object with
iodine renders it still more distinct, and is, therefore, very
important. It is also unnecessary to select parts which
have not completed their growth ; all that is requisite to
obtain the same result is, that they should not have be-
come too hard and dry. I have tried this with many
plants ; but among them I shall only mention the leaves

226 PHYSIOLOGICAL BOTANY.

of Allium porrum, because in the garlics the spherical
clear bodies exist, which exhibit an appearance of one
cell within another, and have sometimes confirmed ob-
servers in the belief that the young cells were contained
within the old ones. We shall call them the secondary
cells. On making longitudinal sections parallel with,
or even perpendicular to, the surface of either the upper
green portion or the under colourless part of the leaf,
and examining them in a drop of water in the usual way
with a sufficient magnifying power, we perceive in the
white part the light cells, which appear clear and trans-
parent ; in the green part, we see here and there a little
of the granulo-cellular matter, which most of the cells
contain, and we also find the light, globular, secondary
cells. But if the sections are moistened for a few minutes
with nitric acid, and then washed with water and coloured
with tincture of iodine, the whole is changed. We now
see, inside the cells, a sac of a yellowish colour, and
almost of the same form as the cells, but more or less irre-
gular, frequently lacerated, more or less detached from
the wall of the latter, and also more or less contracted.
It is completely filled with the granulo-cellular matter,
and when secondary cells are present they are scattered
within the sac, more deeply coloured than the sac of the
cell, and completely filled with granules. The external
membrane of the cell remains transparent, and perfectly
un coloured. But the most remarkable feature exists in the
small warty projections on the margin of the sac, which
fit into apertures in the external membrane of the cell ;
between them this membrane appears raised in roundish
portions, and sometimes we perceive obscurely-defined
laminae in these tumid spots.

After these experiments, I must express my approval
of Hartig's memoir upon the Structure of the Vegetable
Cell. The membrane of the utricle is evidently his
Ptychode, a membrane which descends into the so-called
pores of the external membrane, and is really a distinct
membrane ; it surrounds the internal contents, but

PHYSIOLOGICAL BOTANY. 227

belongs to the secondary layers as it is coloured yellow by
iodine, whilst the outer membrane, Hartig's Eustathe,
and the intermediate layer, Hartig's Astathe, are not
coloured. Hartig must not be offended at my refusing
to adopt these technical terms. They are not only per-
fectly superfluous, but even retard the progress of science;
they form the skins which science must throw off at
every moulting period. The inner membrane of the cell,
or the membrane of the utricle, belongs, with the spiral
vessels, to the secondary formations, and undoubtedly
has some relation to the formation of spiral vessels,
although not that which Hartig has far too positively as-
serted. I shall now immediately pass to

The Life of the Vegetable Cell, together with its Forma-
tion, Growth, Development, and Dissolution. By Dr.
THEODORE HARTIG. Berlin, 1844. 4. This memoir
requires very careful analysis, which cannot be given in
a few words. We shall only make a few remarks upon
it here. In the first section The Life of the Vegetable
Cell, during the period of Cell-formation, the author says,
a, " Origin of cells. Cells are only formed in the inte-
rior of a parent-cell. They are originally simple Pty-
chodal cells with fluid contents, the cell-sap. In the
course of its development, the Ptychode becomes subdi-
vided into an inner and an outer Ptychodal membrane,
in this manner a Ptychodal space is formed, which is
distinct from the cavity of the cell. A fluid resembling
the laticiferous fluid, the Ptychodal sap, is secreted in this
cavity from the sap of the cell. In the Ptychodal sap
the new cell-germ is formed ; this becomes developed into
three different kinds of cells, digestive-, propagating-,
and colour-cells. The digestive- (Metacard-) cells effect
the further elaboration of the sap of the cell. The pro-
pagating- (Epigon-) cells develope a new generation of
cells, of three different kinds, in their Ptychodal cavity,
in the same manner as the original parent-cell. The
colour- (Euchrom-) cells form Euchrome (which includes

228 PHYSIOLOGICAL BOTANY.

Chlorophyll) in their Ptychodal cavity, and starch." Then
follow observations in which, as regards the contents of
the cells, so far as my own investigations enable me to
judge, many of the details are minute and correct. In
fact, too minute, for these contents hardly appear to me
to deserve the name of cells ; the most we could call
them would be cell-nuclei, cell-vesicles, or, with the
author, nuclear corpuscles. They are always of very
different sizes and shapes, never equilateral, even when
closely crowded, hence they are not formed by internal
expansion ; they are never distributed regularly, and often
appear to be perfectly solid internally, like granules of
starch. The secondary cells 3 as I have denominated them
above, are the most regular ; they also contain small cel-
lular granules. The granules of Chlorophyll in succulent
and water-plants are of a tolerably regular figure, but
they appear solid, and altogether of a very different-
nature from the cells which surround them externally.
The cytoblast appears to me to be a granular mass, which
is possibly inclosed by a membrane, but this I will not
decide ; according to the author, it is a perfectly-developed
and not a young cell. He makes the following remarks
upon it : " There can scarcely be any doubt that the cell-
germ of the cytoblast and of the nuclear corpuscles may,
like that of the cavity of the Ptychode, become free and
capable of further development ; but it is equally as cer-
tain that the cell-germ does not originate exclusively from
this source, because it is also formed in exactly the same
manner as within the cytoblast, in other parts of the
cavity of the Ptychode of the cell, where there are no
cytoblasts. In fact, I believe that, as a rule, the cyto-
blast does not produce any propagating cells, but that its
function is rather the elaboration and conversion of the
sap of the cell into that of the Ptychode." If the
author believes the latter to be true, he should not say
that the former can scarcely be doubted. On the con-
trary, it is very doubtful, and has not been proved to
occur by any of his observations. In all these investi-

PHYSIOLOGICAL BOTANY. 229

gations it would be very desirable that the objects should
be carefully distinguished. What applies to the Algae
cannot necessarily be assumed as holding good in the
case of the Phanerogainia, and still less, what is observed
in Fungi, as the author has done. His observations upon
the cells of the ripe and unripe berries of Solatium nigrum
are valuable, but this is a distinct subject, and one which
may be of importance as regards the ripening of the
fruit, and it would have been very desirable for the
author to have instituted a minute comparison in this
point of view. Again, the title, ' The Life of the Vege-
table Cell/ says too much. My friend Hartig knows as
much of the life of the cell as I do, i. e. nothing. Life
is motion, arising from internal excitation, and we are
unacquainted with the movements of the fluids in the
cell which produce development.

Schleiden says, in his ' Principles of Scientific Botany/
p. 200 : " I believe that even in the youngest condition
of the cell, a delicate membrane and a substance which
is not colorable by iodine may be distinguished; the
former of which completely incloses the cytoblasts.
Mohl has apparently (Bot. Zeit., 1844, No. 15 et seq.)
misunderstood me, in relying upon an expression which
was certainly ill-chosen by me, and by which I intended to
illustrate this point, when I first published my discoveries.
But as soon as this primary membrane of the cell has
become even slightly separated from the cytoblast by its
expansion, the whole of its inner surface is very frequently
found covered with a delicate coating of semifluid (very
often circulating in reticularly-anastomosing currents) mu-
cilage, which is sometimes granular, sometimes perfectly
homogeneous and pellucid, but may always be rendered
visible either by nitric acid, alcohol, or iodine; this is Mohl's
primordial utricle." The granulo-cellular mass, called
the cytoblast, certainly always appears surrounded by a
delicate membrane. At first this mass appears compact,
but subsequently it becomes divided, alid then the motion
of the small granules begins to be visible. In the cells

230 PHYSIOLOGICAL BOTANY.

of the pith of the recently- developed twigs of the willow
(e. g.) this mass assumes a tolerably compact form ; in
twigs of a year's growth it is found diffused, and has
formed dotted cells. Now it appears to me, that this
membrane becomes applied whilst in a delicate state to
the wall of the cell, and in certain parts penetrates more
deeply through the secondary deposit, until it reaches
the outermost membrane, and in this manner the appa-
rent holes or pits are produced. The prominences seen
on the utricle after it has been detached by nitric acid,
and which fit into depressions in the membrane of the
cell, and the elevations between them, appear to prove
this to be the case. The nitric acid probably only acts
by contracting the parts and rendering them visible.
The membrane inclosing the granular contents, when
detached from the walls of the cell, is only rendered of a
pale yellow colour by iodine, and at first is not at all
coloured by it. Prom these considerations, it appears
clear to me that this membrane is applied to the outer
membrane of the cells, and that it is again detached from
it by the action of nitric acid, but for this very reason it
cannot be a primordial utricle.

UNGER'S Memoir in the ' Botanisch. Zeit.,' 1844,
s. 498 et seq., upon the Growth of Internodes anato-
mically considered, properly belongs here. The author
counted the internodial cells existing in Campelia Za-
nonia, and then compared the number with their
length and breadth, whence he arrived at the conclu-
sion, that the enlargement of the internodes is the result
of the continuous growth of the new elementary parts,
and also that the enlargement of the internodes of the
axis arises from both the addition of new elementary
parts and the increase of those already existing. He
then goes further, and proposes the question of how, and
in what manner, the addition of new elementary organs
(cells) takes place in the growth of the internodes. He
examined a longitudinal section which passed through
several internodes, and then found that the formation of

PHYSIOLOGICAL BOTANY. 231

new parts occurs in the internodes themselves and not in
the nodes. He moreover says, "If we carefully examine
cellular tissue, in which new formations are in progress,
we find what is very remarkable, that all the cells are not
furnished with walls of equal thickness, but, on the con-
trary, that some of them are delicately constructed, whilst
others are scarcely perceptible. From this we may con-
clude that, in all probability, the latter are of a later for-
mation, and hardly entertain a doubt that any observer
will admit both the fact and the conclusion derived from
it." The next question was, whether the separating wall
was single or double. To determine this point, the
author selected some young hairs which were in the
earliest stage of development, from the recently-formed
leaves of Syringa vulgaris. He endeavoured, by the
action of chemical re-agents, to produce a condensation
and contraction of the finely granular contents, so as to
allow of the more perfect examination of the walls.
Dilute mineral acids answered tolerably, but the result
was best obtained by first subjecting them to the action
of caustic potash, and subsequently iodine. Even then
the separating walls always remain simple. Hence the
author thinks that this forms the commencement of a
subdivision into more cells, and therefore calls this form
of cell-growth merisinatic, but he is too hasty in his con-
clusions on this subject. As Unger expressed his dissent
from Schleiden's theory of cell-formation, the discoverer
of the spermatozoa (or whatever else they may be called)
in the anthers of the Mosses, and the ciliary motions of
the spores of the Algse, &c., receives the following repri-
mand in the 'Principles of Scientific Bot./ p. 210:
' Transverse and longitudinal sections, and a mere glance
through a microscope, be this ever so good, are certainly
not sufficient now-a-days for making phytotomic inves-
tigations."

In the first part of Schleiden and Nageli's ' Zeitschrift
fur wissenschaftliche Botanik' (Zurich, 1844), there is a
Memoir, by Nageli, upon the Nuclei of Cells, Cell-forma-

232 PHYSIOLOGICAL BOTANY.

tion, and Cell-development, wherein Schleiden's theory of
Phaneroganria is brought forward. In the second part
(1845), we find an appendix, entitled, Definition (Beyriff)
of .a cell. The author, after making some remarks upon
Schleiden's definition, says : " The idea of a cell signi-
fies that a portion of the organic matters becomes indi-
vidualised, invested with a membrane, by means of which
it is put in relation externally with the absorption and
exhalation of matters, whilst internally it undergoes
chemical and plastic changes." The commencement is
perfectly correct ; the idea of a cell signifies that a por-
tion of the organic matters becomes individualised, so
that the solid parts form externally an envelope, within
which, in part at least, matters exist in a fluid or even
in a gaseous state. Whether in all cells a solid body is
first formed, does not appertain to the idea, nor has this
been proved to be the case by observation. The nucleus
of the cell, as it appears to me and to others, is an irre-
gular accumulation of granules or vesicles, and seems
more like the mere commencement of a formation, than
a primitive formation itself, which here, as in almost every
other case originates in a liquid. The author says, with
perfect truth, the idea of an organism combines two im-
portant forces that by which it lives, and that by which
it propagates itself. But when it is added, that both of
these depend upon its being composed of cells, this
dependence certainly is not apparent. If it is proved
that the motion of Browne's molecules depends upon
an internal influence, they are alive, whatever may
be their internal structure. The organism requires a
reciprocal action of the parts upon each other as organs,
which is certainly most easily produced by the movement
of fluids within them ; but it by no means follows, it is
not proved, nay, it is opposed to experience, that it is
entirely composed of cells. If it were stated that it is
produced from cells, my reply would be, that this is very
probable, but nothing more.

In the search for definition of a plant and of the
vegetable kingdom, much dependence is placed upon the

PHYSIOLOGICAL BOTANY. 233

absence of nitrogen in the membrane of the cells of plants,
and also upon the presence of nitrogen in the animal
kingdom. But supposing that nitrogen were to occur in
the membrane of several vegetable cells, would the plants
for this reason, cease to be plants? Boussingault has
shown that a large quantity of nitrogen exists in plants,
but where, has not been positively determined. It is
reversing the proper method to commence with chemistry
in the study of natural history : and this, first, because
chemical analysis is the most difficult ; secondly, because
its resources are inexhaustible, and cannot be regarded
in the same light as the latter ; and lastly, because it
affords us no insight into the intimate structure of organic
bodies, as is proved by isomeric substances. The mem-
brane of plants is isomeric with starch, as Pay en has
shown, and yet the two are very different from each
other.

Investigations upon the Growth of Cells. By Dr.
SCHAFFNER of Herstein. Flora, 1845, 451. "If we may
be allowed/' says the author, " to deduce conclusions
from the investigations which have been detailed, the fol-
lowing cells increase by primary cell-formation. 1. The
cambium cells (which are subsequently developed into
prosenchyma and vascular cells). 2. The cells of the
liber, which do not differ materially in their earliest stage
from those of the cambium, but form a distinct system.
3. Part of the cells of the parenchyma, to which the cells
of the leaf (excepting those of the cotyledons), and the
cells of the parenchyma of the fruit of the apple and the
plum provisionally belong, i. e. should the absence of
secondary cells in them be confirmed." This is of no im-
portance, the point is whether the so-called secondary cells
are really such, i. e. are formed from the parent-cell. " By
the formation of secondary cells, the remaining cells of the
parenchyma increase, as seen, e. g. in the cells of the pith
and of the bark, &c." (???) " Increase of cells by division
certainly does not occur in Phanerogamous plants." (?)

234 PHYSIOLOGICAL BOTANY.

In an appendix upon the laticiferous vessels, he advises us
not to read upon the subject at all, because most of what
is stated upon it involves much contradiction ; he quotes
Bischoff and Schleiden. He has not mentioned my re-
searches. In the first part of my Lectures on Botany, he
would have found much upon this subject ; and also many
figures in the Plates to the first part of my Anatomy of
Plants.

Researches on the earliest Modifications of Organic
Matter and the Formation of Cells. By M. Coste.
Comptes rendus, 1845, 2, 911, 1396. This memoir
principally treats of the animal cell, and tends more by
reasoning than experiments to show that the theory of
the production of cells by the nucleus is not based upon
satisfactory investigations.

The preceding experiments upon the manner in which
new cells are formed, induced me to make some extended
observations myself. Thus, when our object is to become
acquainted with the manner in which cells are reproduced
in the Phanerogamia, without confusing the phenomenon
with others not appertaining to it, the method of proceed-
ing used by linger is the best. With this view I caused
some bulbs of Allium cepa to grow in a hyacinth glass
filled with water, and made marks upon the roots which
had grown out, with Indian ink, one close to the bulb,
another close to the conical apex, and a third midway
between the two former. In a few days the roots had
grown very considerably, the conical apex not so, as far
as was distinguishable, the base had grown but little ;
the portion between the apex and the middle had grown
most. The latter portion was again divided in the middle,
and it was found that the part nearest to the apex had
again grown very considerably, whilst that next the middle
had grown but little. On making a longitudinal section
from the mark at the apex to near the upper mark, treat-
ing it with nitric acid, and then iodine, a great many short
cells were seen near the apex, these were found gradually to

PHYSIOLOGICAL BOTANY. 235

increase in length towards the upper part, and finally be-
came very long. But the cells at the circumference of the
root were longer than those in the centre. In all of them
the inner membrane was separated from the walls of the
cells, and was contracted around the granular contents,
which were of a dark brown colour. The sac thus formed
assumed the shape of the surrounding cells, the walls of
which did not appear at all coloured by the iodine. Each
cell contained the globular sac, which I denominated the
secondary cell above ; it was also of a brown colour, and
filled with a granular mass. It was always situated in
the longer sac, but in different cells sometimes at the
ends, sometimes in the centre, sometimes near the centre.
Thus, where the growth was most active, short cells ap-
peared to have been formed, which then became elongated
and had completed their growth. I could not satisfac-
torily perceive any division of cells. I had made similar
marks to those upon the roots, on the young leaves which
sprouted out from the same bulb ; one near the bulb,
another just beneath the apex, and a third midway
between the two other marks. That nearest the apex
remained unchanged ; the apex of the leaf, just as the
apex of the root, did not grow ; that portion between the
mark on the apex and the middle had increased but little,
just the reverse of what occurred in the roots, in which
this was the part which had grown most, whilst that part
of the leaf near the base had increased very considerably,
whilst in the roots, on the contrary, it had grown but little.
A longitudinal section of a marked leaf was then made
parallel with its surface, from the base of the leaf at the
root upwards, and treated as before. In this case, as had
previously been found in that of the root, short but not
broader cells were found at the base of the leaf near the
root, where the growth commenced, and these increased
in length upwards, towards the middle of the leaf. The
formation of these short cells and their elongation evidently
produces the increase in size of the leaf, as I have already
remarked in my < Lectures on Botany/ (p. 83,) and have

236 PHYSIOLOGICAL BOTANY.

had figured in my Plates (part I, pi. i, fig. 4/5). There
was no appearance of the formation of one cell (secondary
cell) from another (parent-cell), and the inner sac remained
unaltered, still forming an inner sac, and certainly not
becoming external. This is what ensues during the
growth of the cells in the parts of the Phanerogamia.
The phenomena which occur in the embryo- sac, or in the
cells of the Algae, which, as the remarkable phenomenon
seen in the cells of the Spirogyra show, perform different
functions from the cells of the Phanerogamia, do not
belong here, and no conclusion can be drawn from these
so-called cells as regards the cell properly so called.
The apices of the roots and leaves which do not grow,
consist of very short cells, all of which contain a nucleus
of considerable size ; this, however, as in the other cells
of the leaf and root, never becomes developed into a
distinct cell.

On the Penetration of the Cuticle into the Stomata. By
H. v. MOHL. Bot. Zeit., 1845, p. 1, and Ann. and Mag.
of Nat. History, vol. xv, p. 217. The different state-
ments which have been made upon this subject induced
the author to institute some investigations upon it. For
this purpose he adopted the method of soaking the
sections of the leaves for examination in tincture of
iodine, washing them with water, and then submitting
them to the action of sulphuric acid. The latter not
only heightens the yellow tint of the cuticle when coloured
by iodine, but it has especially this advantage, that the
cells of the epidermis of most plants are disintegrated,
with the production of a blue colour, or entirely dissolved,
according to the strength of the acid employed, and the
cuticle can then be very readily distinguished and separated
from them. The general result obtained from these in-
vestigations was, that, as asserted by Payen, a direct
prolongation of the cuticle penetrates into the stomata,
and runs down between the cells bordering the orifice to
the air-cavity, in the form of a tube very highly com-

PHYSIOLOGICAL BOTANY. 237

pressed on both sides. In the opinion of the author, no
doubt can be entertained, after careful examination, that
this tube is not closed either at the entrance into the
stomata, or lower down, between the cells of the orifice.
When it has arrived at the inner termination of the
stomatic aperture, this tube dilates into a smaller or
larger funnel-shaped expansion, which clothes the inferior
surface of the epidermis so far as this bounds the air-
cavity externally. This funnel-shaped expansion presents
some varieties in different plants, which the author details.
Thus the cuticle either lines the walls of the air-cavity
only, without penetrating into the intercellular passages,
or it penetrates into some, or even all, of those passages
which are in connexion with the air-cavity. Lastly, the
author comments upon the question of the cuticle being
a peculiar membrane, different from the epidermis. He
believes that it is not so, but that its peculiarity arises
from a change in the substance of the external layers
of the epidermal cells themselves. Were I to pass
hastily over this paper, as the author says is my custom
(although not very politely), I should say that the point
is not how the cuticle is formed, but whether it is com-
posed of the outer walls of the epidermal cells, and as
this has not been shown to be the case, it must be con-
sidered as a peculiar membrane until it is so. At all
events, the question remains in statu quo. But, as in all
questions relating to the formation of organic bodies, we
must look forward for a complete elucidation of this point,
to a clearer insight than has yet existed.

Investigations upon the Cellular Structures which Jill
up Vessels. By an anonymous Author. Botan. Zeit.,
1845, p. 225. The author first shows that these bodies
consist of true cells, or, as he expresses it, that they are
phenomena analogous to the ordinary simple cells of
plants. These cells are not generally formed while the
plant is young ; in first year's shoots of Vitis vinifera and
Sambucus niyra, as also in the stems of Cucurbita pepo,

238 PHYSIOLOGICAL BOTANY.

the vessels were empty in summer ; towards the end of
October and at the beginning of November they only
contained a small number of cellules adhering to the walls
of the vessels ; but a month later he found them copiously
furnished with both large and small cells . In a fourth
year's branch of Robinia pseudacacia, the most external
annual zone was in a condition similar to that of a first
year's shoot of the same plant ; the three inner ones were
completely filled with cells. As regards their being
adherent, he makes the remarkable observation, that the
small cells are always attached to the side of the vessel
when it is surrounded by cells of woody tissue or the
parenchyma of the medullary rays, but never to a wall
which is bounded by an adjacent vessel. Moreover he
observed that one of these cells always lies in front of
the dot of a vessel, which corresponds with the dot of the
adjacent external cell. He also believes he has observed
that the membrane of the utricle has some relation with
the primary membrane (which belongs to the external
cell and the vessel, and which closes the two dotted canals),
and that in its earliest stage it is an expansion of this
primary membrane into the cavity of the vessel. Hence
the formation of the inner cell depends upon the activity
and development of an external adjacent cell. This is
most distinctly seen when preparations of these vessels in
Vitis vinifera and Sambucus nigra are treated with
caustic potash. To avoid unnecessary circumlocution, he
adds, he cannot avoid assigning names to the objects,
especially to distinguish the old cell from the utricular
sac, which he denominates the tliylle, these two composing
one compound organ. Considerations upon the forma-
tion and development of these and other cells follow.
The investigations of the author deserve the greatest
attention and careful repetition, for the purpose of either
confirming or correcting these observations upon a re-
markable phenomenon.

We have received accurate and comparative experiments

PHYSIOLOGICAL BOTANY. 239

on the chemical peculiarities of the vegetable cell, first from
Payer), he having previously made his excellent observa-
tions on Starch. The whole of these investigations were
published in 1842, in his ' Memoires sur les Developpe-
mens des Vegetaux/ He first instituted his experiments
upon the cellular tissue, which consists of little more than
membrane; this was taken from very young parts, e. g. the
ovule of the almond, pear, and apple tree, and of Heli-
anthus annum ; the delicate membranes formed upon the
coagulated drops of fluid which exude from the sections
of the vessels of the cucumber, also upon the pith of
young shoots of Samhucus nigra, cotton-wool after a first
and second purification, and the spongioles of roots, and
the pith of Aescliynomene paludosa (rice-paper). All these
substances were repeatedly treated with dilute hydro-
chloric acid and ammonia, washed with water before each
repetition of the process, and lastly exhausted with
alcohol and ether. They were then strongly dried, pow-
dered as far as possible, and then burnt with oxide of
copper. He found, as the result of the elementary analysis,
the composition C 24 H 23 O 3 , which is isomeric with starch.
He then gives a simple, direct experiment, by which
cellular membrane may be recognised under the mi-
croscope. A small section, e. g. of rice-paper, is placed
under the microscope in a drop of water; one or two
drops of an aqueous solution of iodine are then added,
which produce a pale-yellow colour ; and lastly, a drop of
concentrated sulphuric acid. The membrane is then first
coloured blue, and is finally wholly dissolved, so that
only yellow traces of the matters which were contained in
the membrane remain. A better method than this one
of Payen's, is to place the section for microscopic exami-
nation in a drop of water, then to add a drop of nitric or
hydrochloric acid ; to let the whole remain for about two
minutes, then to wash the section with water, and to
colour it with tincture of iodine. The membrane itself
now appears perfectly colourless ; sometimes it is slightly
bluish here and there, from a portion of starch which is
dissolved, and all the foreign matters are rendered deep

240 PHYSIOLOGICAL BOTANY.

yellow, so that they are readily distinguishable from
the membrane. The experiments upon the leaves of
Allium porrum and the roots of Allium cepa, which have
been already detailed, were made by this method. It
must be remembered that the starch-granules are here
dissolved ; hence, if it be required to observe them, no
acid must be used. If this experiment be reversed, and
the section be first examined with tincture of iodine, the
starch is easily recognised ; but no nitric acid must be
added, because it dissolves the iodised substances, leaving
the membrane, which cannot now, at least very readily,
be recognised and sketched. But by this method the
membranes of the utricles, the contents of which are
dissolved, are distinctly seen within the cells. Caustic
potash and caustic soda also remove the contents of the
membranes, and leave the latter, although in an indistinct
state. But I must return to Payen's investigations. He
next examined, by elementary analysis, the following
structures, after having exhausted them with several
solvent media. The leaves of Endive and Ailantlms
glandulosa, the internal cellular tissue of Agave Ameri-
cana, the spiral vessels of Musa Sapientum, the radicles
of maize; portions which had resisted the digestive
process in animals, the albuminous tissue of maize and
corn, the albumen of Phytelephas, and the kernels of the
date ; the hairs of the seeds of the Virginian poplar, the
vegetable membranes of which the nest of the wasp is
constructed ; the heart- wood of the oak, the wood of the
Coniferae, also Conferva rivularis and oscillatoria, the
membrane of Agaricus edulis, probably Ag. campestris L.
He also makes use of the name Scariola, as well as
" Chicoree endivie" How can any chemical investiga-
tions be of use, if the object which has been examined is
not definitely stated? Then follow investigations upon
the substances existing in the cells which also contain
nitrogen. I have quoted these details here for the pur-
pose of introducing a memoir by Fromdery, on Cellulose,
given in the ' Scheikundigen Onderzoekingen,' 2 D. s.
36, and which is extracted into the ' Journ. f. praktische

PHYSIOLOGICAL BOTANY. 241

Chemie/ 32, Bd. s. 198. He subjected Cetraria Islandica
and Ayaricus albus to elementary analysis, and his results
agree tolerably well with those of Pay en. He then makes
the following remark : " I am also satisfied as to the
perfect accuracy of his experiments, yet I cannot deny that
I am astonished, first, at not finding it stated* anywhere
that he previously determined the amount of ash, except
in his first memoir (Annal. des Sciences Natur., 2 ser.
t. ii, p. 27), since even if he had not found any ash pre-
sent, he ought to have mentioned it; moreover, since
none of the substances mentioned by myself as having
been analysed were perfectly free from the so-called in-
crusting matter, and since the results of Payen lead to
the same conclusion, which is explained by the intimacy
with which these substances permeate the primary cellular
tissue ; and again, since silica, which is so generally dif-
fused throughout the vegetable kingdom, would very
probably have entered into the composition of these
matters, it does not appear possible that the vegetable
structures subjected to examination could have been per-
fectly free from silica." This suggestion is quite correct.
Payen gives the amount of ash contained in the vegetable
structures when not yet separated from the matters depo-
sited upon the cellulose. We thus find that 10*80 p. c. of
silica are stated to exist in the leaves of Endive, but none
in the leaves of the same plant when exhausted of every-
thing but the cellulose. This is very improbable, for the
amount of silica in the leaves of the Graminaceas, before
purification, is stated to be 12'25 ; but I find no analysis
given of the leaves after purification. But in this case
the amount of silica existing in the cellulose must be very
large, for the incinerated leaf is so completely converted
into silica, that all its parts can be accurately distinguished
under the microscope ; a remarkable phenomenon, and
one which still requires careful investigation, because it
is opposed to what we know regarding cellulose.

In the same ' Scheikundige Onderzoekingen/ 1. c. p. 62,
' Journal f. Prakt. Chemie,' 1. c. pu 204, we have an

242 PHYSIOLOGICAL BOTANY.

Analysis of the Seeds of Phytelephas, Ruiz et Pavon
(Elephantusia Willd.),* by BAUMHAUER. He gives his
results in the following words : " Our results show dis-
tinctly that the perisperm of Phytelephas does not, as
Payen states, consist of pure cellulose, contaminated with
albumen, two nitrogenous substances, silica, two fatty
bodies, and salts ; but that, in addition to these matters,
of which the albumen, the two nitrogenous substances,
and the two fats are in extremely small quantity, an ad-
ditional deposited substance occurs, which differs but very
slightly in its per centage composition from cellulose."

We shall take this opportunity of subjoining the ob-
servations which have been made upon Starch during this
period. First :

Description and Figures of some remarkable Forms of
Granules of Starch in the Root of Sarsaparilla, and in the
Rhizome of Hedycliium Gardnerianum. By G. BISCHOFF,
Bot. Zeit., 1844, p. 385. The granules in the former
root are very often of a hemispherical or half-ellipsoidal
form, moreover they are united by their bases, or there
are four or more granules connected together in a regular
manner. These various forms are accurately described
and figured. The author compares them to the combi-
nations presented by many kinds of pollen-granules, they
might also be compared with a tricoccous or tetracoccous
capsule. Several others might be added. I have met
with a form in which a small angular grain occupied the
centre, and the other five were so arranged around this,
that the whole figure somewhat resembled a regular pen-
tapetalous flower. The author remarks that the concentric
lamination was not perceptible in daylight, but it was
distinctly so by subdued lamp-light. The author also
found potato-starch to consist of a combination of two
granules. The starch existing in the rhizomes of the

* One botanist considers that the alteration of the word Phytelephas into
Elephantusia would be a very unnecessary change. ^i Phytelephas signifies
a vegetable elephant, and siich a zoophyte is something too terrible.

PHYSIOLOGICAL BOTANY. 243

Scitaminese is singular enough. The granules are cylin-
drical, and either curved or bent at an angle ; they are
sometimes club-shaped, or of various other forms, some
of which resemble a pileate Fungus, and in consequence
of their being contracted between the rings, they distinctly
exhibit their laminated or tunicated structure, each main-
ring again exhibiting a larger or smaller number of ex-
ceedingly delicate, parallel, curved, transverse markings.
The larger segments undoubtedly denote the separate
granules of the composite structure, and each of these is
again finely laminated.

On the Starch of Gloriosa Superba, L. By JULIUS
HUNTER. Bot. Zeit., 1845, p. 193. The form of the
granules of starch which exist in the rhizome of this plant,
is sometimes that of a perfect sphere or ellipse ; but by
far the larger number of granules are bounded by one or
more plane surfaces, which sometimes meet at an acute,
sometimes at a right angle. If we divide an egg, says
the author, through its middle, at right angles to its
long axis, we obtain two kettledrum-shaped halves, these
accurately represent, on a large scale, the appearance
frequently exhibited by the starch of Gloriosa. Other
pieces resemble that form which we should obtain on
dividing an egg anywhere parallel to its longitudinal axis;
other forms again represent sections of a sphere, i. e.
pieces which are bounded by two plane surfaces inter-
secting each other at an angle of 120 and one spherical
surface. Sometimes we find three plane and one spherical
surface ; and lastly, we have also pure stereometric forms,
pentahedra, hexahedra, and octahedra. Occasionally the
granules are of an indefinite form, which does not admit
of description. Maranta bicolor Kerr, and Jatropha
manihot, also exhibit pentahedral granules of starch. The
author brings these forward as a proof that even an or-
ganic compound may assume a crystalline form, and from
this consideration he applies to them the term glandules.
It is then found that these glandules become disintegrated
when removed from the cell and placed in water, upon

244 PHYSIOLOGICAL BOTANY.

the object holder, which is not usually the case, for in
other plants the granules retain their connexion. The
author then passes to the investigation of how these
granules of starch are formed and developed. We might
at first imagine, says he, that as in crystallization, the
amylaceous plasma (the analogue of the saline solution)
is deposited upon the little globules which first separate,
and that in this manner, by the continuation of depo-
sition upon their outer surface, larger granules are formed.
We might add to the author's explanation that the
granules are formed around a nucleus, as is usually con-
sidered to be the case from the manner in which crystals
are formed. According to the author's view, the twin
granules of the nucleus of the one individual must lie
close beside the nucleus of the other individual, and
nearly in that plane in which both are connected, or
near the parchment of the kettledrum, if the above
comparison is retained. This is not the case, for the
nucleus is situated in the bottom of the kettledrum, at
the end of the elliptical or spheroidal section (this is
also shown by the figures given by Bischoff.) The
author then proceeds to the question whether the pres-
sure of x the cell inclosing them might not produce an
angular form. But this is not the case, for the granules
have an angular form, even when they do not completely
fill the cells. It is evident, says the author, from what
we have stated, that pressure cannot be the cause of the
formation of the gland-like collection of starch. From
all this, he adds, all that remains is the prospect of a
peculiar formative process in the vegetable kingdom. Re-
garding the vegetable cell, we know with perfect certainty
that the concentric appearances, as seen e. g. in the stony
tissue, as it is called, of the pear, &c. arise solely from
the centripetal formation of layers. But nothing is op-
posed to the view that the layers of the granules of starch
also are formed by centripetal, i. e. internal deposition ; on
the other hand, this hypothesis is supported by the fact
that the nucleus, as it is called, of Fritzsche, or the central
cavity of Schleiden, contains much water, and is, as it

PHYSIOLOGICAL BOTANY. 245

were, gelatinous. For as soon as sulphuric acid is added to
the granules of starch, under the microscope, and it begins
to remove the water from the inner layers, a bubble of air
appears in the place of the nucleus ; the same occurs when
the granule of starch is heated ; in fact, even when fresh
starch is dried at the ordinary temperature of the air.
The latter phenomenon, which was not observed by either
Schleiden or Fritzsche, thus also explains the formation of
the fissure near the nucleus. But if, as appears from
these observations to be the case, the nucleus and the
layers nearest to it, contain more water than the outer
ones, i. e. if they are softer and less consolidated than the
outer layers, we might assume with tolerable certainty
that the central layers which surround the nucleus are the
youngest, and the peripheral ones the oldest. Now, if
this hypothesis be retained as the most probable, there is
no difficulty in explaining the spot where the nucleus
should be found. Accordingly as the layers happen to
be thick or thin, the nucleus must be situated more or less
excentrically, in fact it must be excentric in the large
globules. For as soon as the centripetal formation of
the layers was uniform in all parts of the inner surface, a
condition would occur, which would prevent any further
development, because the walls being everywhere of equal
thickness, the transmission of new nutritive matter would
be prevented, whilst this condition would never occur if one
part of the granule were thinner than the rest. When
the walls of the cells are thicker, other means come into
play in facilitating the access of nutriment, viz. pore-
canals. The author moreover adds : " We must rest
satisfied for the present, with the conclusion which has
been arrived at negatively, that a process similar to that
of cell-formation must be admitted to occur in the case
of the granules of starch, the nature of which must form
an object of future investigation." It is very pleasing to
find that the author has differed from the ordinary ex-
planations of the formation of starch. I quite agree in
his opinion, that the granules of starch are formed from

246 PHYSIOLOGICAL BOTANY.

without inwards, and that this is produced by a peculiar
process of formation, which certainly resembles the process
of cell-formation, but is not always perfectly regular.
I should attribute the eccentricity of the layers around
nucleus, solely to this irregular formation. The granule
of starch apparently absorbs moisture on all sides, and
then developes the layers internally. A similar internal
formation is also the cause of the regular separation of
the granules in the root of Sarsaparilla, which then finally
passes into the external crystalline form of the granules
in the tubers of Gloriosa superda, as was first discovered
by the author. All the granules existing in the same
tuber, even those close together, are not of the same form ;
some are more rounded externally, some are rounded on
the sides, bounded on the others by two planes, because
originally they separated from each other at that part,
others are bounded on all sides by plane surfaces, like the
central grain in the compound form, presented by the
granules in- the root of Sarsaparilla. I should always
ascribe these crystalline forms to the internal separation
of granules, to which view I was led by the granules of
starch in the bulb of Ornithogalum (MyogalumJ nutans.
But the author will publish his own investigations upon
that point. He then adds some remarks upon Schleiden's
observations on this subject in his ' Systematic Botany.'
The amorphous granules of the seeds of Coriandrum minus
arise from desiccation ; this is also the case with the cup-
shaped granules of starch in the rhizome of Iris pallida.
Schleiden, in opposition to Meyen, incorrectly denies the
occurrence of discoidal granules in the Cannaceae ; for in
Canna variabilis for example, we find only such. The
arrow-root of commerce presents considerable differences ;
the author details them. Most of the commercial article
is obtained from Tacca pinnatifida ; the same statement
applies to sago. He could not detect any cup-shaped
granules in Ead. (stolones) Iivarancuste, such as Schleiden
states to occur. I look forward to the continuation of the
author's accurate and valuable investigations.

PHYSIOLOGICAL BOTANY. 247

Observations on the Formation of Starch. By C.
MULLER; Bot. Zeit., 45, 833. They were instituted
upon Chora crinita, and, according to the author, they
exhibit the following points : it is the cytoblasts which
are transformed into starch, and this only occurs in cells
which are mature.

Note upon the Phenomena produced by the Transmission
of Polarized Light through Starch. ByM. BIOT. Compt.
rend., 1844, vol. i, p. 795. In his previous experiments
the author examined the granules of starch by means of
two prisms, one of which was placed above the other, and
crossing it at right angles ; on the present occasion he
changed the apparatus by so placing a plate of mica be-
tween the two prisms, that the median line between the
two axes formed an angle of 45 with the principal
sections of the prisms. The mass of the granule is then
seen illuminated with bright colours, the tints of which
vary with every change of position, and with the direction
in which the luminous rays are transmitted ; so that, as
in a picture, all the curves of the outline, all the undu-
lations of the surface, all the peculiarities of the structure,
and the slightest accidental alterations become sensible.
Probably under some circumstances this means may prove
of the utmost value ; but in the present case, considering
the great and accidental varieties in the structure of the
granules in starch, it is perhaps of minor importance.

We shall now pass from cells to vessels. In the
' Annual Report/ for 1841, I made some observations
upon the work of C. H. Schultz, on Cyclosis in plants,
author has replied to them in a book which we shall

lude to presently, viz. : ' Discovery of the True Nutri-
tive Process of Plants/ Berlin, 1844. He there says,
with regard to my remarks, p. 54 : " There are two main
points to be considered : first, whether I have correctly
denominated the vessels of the liber, vessels of the vital
fluid (lebenssaftyefassc] ; and, secondly, whether the cur-
rents of fluid in Commelina codcstis consist of a circula-

248 PHYSIOLOGICAL BOTANY.

tion (rotation) of the granules without any trace of anasto-
mosis, as in Vallisneria." He has distorted my statement,
and I must therefore briefly repeat my remarks. Schultz
was the first to detect the motion of the fluids in the
so-called proper vessels (vasa proprid], he also first gave
good illustrations of these vessels. But to carry out his
hypothesis regarding Cyclosis, he has attributed these
vessels, which he denominates vessels of the vital fluid
(laticiferous vessels), to many plants in which they do not
exist. They are said to occur in the bark of many trees,
as the birch, but I only find vessels of the liber in it,
and no one has seen them in it, even the author himself
only represents their transverse not their longitudinal
section, thus we are ignorant as to whether he has really
seen them or not. This is most striking in Commelina
ccelestis, in the stem of which plant, near the spiral vessels,
ramified laticiferous vessels are said to pass out and
spread over the adjacent cells. He has given a figure of
this. But I find, besides the spiral vessels, merely rows
of pareuchymatous cells, in which granules circulate as
in the cells of Vallisneria ; next to them, are more rows
of broad cells, in which currents of fluid are visible, but
they are certainly not inclosed in vessels. Therefore
there is no trace of ramified vessels. The result is, that
the motion of the fluid in the so-called laticiferous vessels
is the same kind of motion as had been previously found
in plants, namely, as the circulation in the cells of plants,
discovered by Corti, which was first accurately observed
by Meyen in Vallisneria., and the currents of fluid which
Robert Brown first observed in the hairs of Tradescantia.
The motion of the liquid in the articulations of Char a is
of the same kind. The author makes the following re-
mark among others : "It is to be regretted that the
author should have been able to gain so little by the
most earnest exertions and sacrifices to solve this pro-
blem, that he has rather misunderstood it altogether,
and by useless opposition to the evolution of truths, the
great importance of which was first recognised abroad,
deprived himself the renown of having aided in their

PHYSIOLOGICAL BOTANY. 249

promotion." An example of the incomprehensible arro-
gance of the author who is bound to a fixed idea. The
Academy of Paris certainly awarded him the prize for
his Memoir upon the proper vessels, for which no one
will censure them, but they declared at the same time,
that they do not participate in his views. No academy
can test the accuracy of the individual details which are
contained in the prize-essays, nor could the Paris academy
do so in the present instance.

There is an analysis of the milky juice of Asclepias
Syriaca, by the same author, in the ' Flora/ 1844, p. 374.

Phytological Studies. By M. le Comte de TRISTAN.
4th Mem. Researches upon the Laticiferous Reservoirs
and Canals, Ann. des Sc. Nat., 38, t. i, p. 176. This
memoir is directly opposed to Schultz's views. Some
parts of plants do not contain any laticiferous vessels,
hence they cannot serve for nutrition. On the properties
of the latex. Varieties in the laticiferous vessels. It is
impossible to condense this memoir into the form of an
abstract.

STEM AND ROOT.

On the Dependence of the Increase of Thickness of
Dicotyledonous Trees, on the Physiological Activity of the
Leaves. By H. MOPIL. Bot. Zeit., 1844, p. 89. Ac-
cording to the theory of Dupetit-Thouars, says the
author, the lateral growth of the stem is in connexion
with the expansion of the buds, therefore with the for-
mation and development of new leaves, and depends upon
the circumstance that the buds, in the same manner as a
germinating plant, send out radical fibres which descend
between the bark and the stem, and form a new layer of
wood ; according to another theory, the lateral growth of
the stem depends upon the activity of the leaves, because
they prepare the nutritive fluid, which is applied to the
formation of new woody layers. To decide this point,

250 PHYSIOLOGICAL BOTANY.

the author measured the circumference of some trees,
which were about eight years old and in an active state
of growth,, at various periods from the commencement to
the end of the vegetating season, and calculated the
mean daily increase in the circumference of the stem for
each of these intervals of time. The trees were, Gymno-
cladus Canadensis, Gleditschia macracantlia, Tilia argentea,
Populus Grceca, Pavia lutea, and Morus alba. A table
of the increments is added. The following details are
selected from the author's remarks upon his experiments :
On the 22d of June the terminal buds of Pavia lutca
were already formed, but the lateral growth, instead of
ceasing, from the above period to the 22d of August,
somewhat increased; it then diminished to a small amount.
From the 2d of March to the 22d of June, hence before
the development of the terminal buds, the circumference
of the stem increased 11 '8 millimeters ; from the 22d of
June to the end of the year 16*2 millimeters; so that
by far the greater proportion of the increase occurred
during that period in which no leaves were developed.
The same was found to be the case, although less
strikingly, in Gleditschia and Gymnocladus. Hence the
author draws the conclusion, that these observations are
directly opposed to the theory of Petit-Thouars. Valuable
as they are in themselves, the adherents of Petit-Thouars
will not rest satisfied with them, but will object that the
roots of the buds, between the bark and the stem, which
cause the lateral expansion of the stem, would be small
and delicate at the commencement, but that they increased
with the activity of the leaves, and in this manner aug-
mented the thickness of the stem. The author adds,
that the circumference of the stem also increased, although
to a small extent only, when the buds began to enlarge
and expand. Hence he thinks that nutritive matter,
prepared the preceding year, is applied to the first ex-
pansion of the stem in the spring, without the leaves
preparing it. Why not ? Although it appears determined
by numerous experiments, that the leaves serve for the

PHYSIOLOGICAL BOTANY. 251

preparation of the nutritive fluid, there is at present no
ground for limiting this preparation to the influence of
the leaves alone, if the observations definitely indicate
another mode. Lastly, against Agardh's view, that trees
grow principally in length during the first half of the
summer, but principally in thickness during the second :
which is not in conformity with observation.

On the Growth of Internodes, considered Anatomically.
By Professor UNGER. Botan. Zeit., 1844, p. 489.
This memoir has already been spoken of at p. 230. It was
necessarily brought forward there in connexion with the
origin of new cells by division, a mode of formation which
I should confine to the Algas alone. Here we have to
notice the general growth of the parts, of which the author
very modestly says, that in one special instance i. e.
Campelia Zanonia it takes place not only by the forma-
tion of new cells, but also by the increase of those
already formed. This law might, in truth, be extended
to all the Phanerogamia at least.

In the ' Coniptes rendus/ 1844, t.i, pp. 899 and 972,
we have the fourth notes relative to the protest of M. C.
Gaudichaud, of which we have already spoken.

Continuation of the Anatomical and Physiological
Researches upon some Monocotyledons. By M. MIRBEL.
(Second Memoir.) Compt. rend., 1844, ii, 689. In
this memoir the author gives a very accurate description
of the stem of Dracaena Australia (Cordyline Australis),
as regards its internal structure, and especially the course
of the vascular bundles. He endeavours to prove that
the latter arise from the root, and from the inner wall of
the stem. He has not only described the stem when
perfect, but also when young, and this with great care.
I first expressed my views on this subject to the Congress
of Italian philosophers at Milan, and they are published
in the ' Atti della sesta reunione degli Scienziati Italiani

252 PHYSIOLOGICAL BOTANY.

tenuta in Milano/ Milan, 1845, vol. iv, p. 511 ; more in
detail in the ' Flora/ 1845, p. 272 ; also in rny ' Lectures
on Botany/ part ii, Berlin, 1845, p. 309. The Date-
palm is the subject there treated of. In germination,
the embryo or the cotyledon becomes elongated, as is
usual in the Monocotyledons, and splits into a sheath,
from the base of which the stem grows upwards and the
root downwards. The former, which is surrounded by
the sheath, contains within it a small tuberous body,
consisting of parenchyma and delicate spiral vessels
passing round it; above, it immediately forms a bud,
consisting of leaves only, as in the Monocotyledons
generally. The leaves acquire considerable length, whilst
the stem remains as an almost spherical tube. If it be
examined at the end of several years, about six or eight,
its section exhibits a nucleus, which is throughout tra-
versed by a plexus of vascular bundles, which interlace
in every direction. The nucleus is surrounded by a
cortex of parenchyma; at the upper part, beneath the
bud, there is also a layer of parenchyma forming a cortex,
through which vascular bundles pass from the nucleus to
the leaves. Thus the young Palm perfectly resembles a
corm, which only differs from a true bulb in the absence
of the fleshy scales. On making a section of the tall
part of the stem of a Date-palm, we find a number of
vascular bundles traversing it longitudinally. The nearer
they are to the circumference, the closer they are toge-
ther; and at the very circumference they are most
closely crowded, whilst towards the centre they are more
scattered, more surrounded with cellular tissue, and they
are most scattered at the very centre. On closely examining
the woody bundles, we find that they are by no means
parallel to each other, but interlace in various ways,
forming, however, but very small angles with each other.
Hence the stern of a Palm is a longitudinally extended
corm.

Neither Mirbel nor Gaudichaud have noticed this bulb-
like condition of the young Palm-stem, nor have they

PHYSIOLOGICAL BOTANY. 253

definitely alluded to the fact that the Palm grows at the
summit only, and that the vascular bundles arise from
the interior solely at that part, and grow towards the
leaves. Hence I cannot agree with Gaudichaud's opinion,
that the vascular bundles arise from the leaves, although
he has often quoted me amongst a singularly-enough se-
lected series of authors who are of his opinion. Neither,
on the other hand, can I agree with Mirbel, in considering
that vascular bundles arise from the interior of the stem.
The growth takes place at the summit only, and there
the vascular bundles come out from the interior.

Gaudichaud's "Memoirs against Mirbel," are in the
'Compt. rend./ 1845, i, 1375, 1436, and 1677, and
ii, 99, 20], and 261; the memoir upon the Stem of
Ravenala, in the same year (ii, 391), must also be
referred here.

Dr. v. Martius has written a paper Upon the Process
of Growth in Palms, especially as regards the Course of
the Fibres in the 8tem, which was published in the
' Gelehrte Anzeige' of the Royal Academy of Sciences of
Bavaria, for February, 1845. The author communicates
his results to the Academy of Sciences at Paris ; hence
they are given in the 'Compt. rend./ 1845, i, 1038.
Gaudichaud has expressed himself strongly upon and
against them (p. 1207). Mirbel also is dissatisfied with
them. This was to be expected; he who takes the
middle path is repulsed on both sides. The vascular
bundles, says the author, originate at the summit of the
growing point, in the nucleus of the bud or " pJiyllo-
phore" (according to Mirbel), in the substance of the
more recent cellular tissue which is in course of develop-
ment, and which here forms a peculiar layer, coating the
older and subjacent parts like a mantle ; the recent ones
being always formed externally to, and more or less above,
those already developed. This is explained in the sub-
sequent parts of the memoir as follows : As the young
plant, even in its first stage after germination, is furnished

254 PHYSIOLOGICAL BOTANY.

with conical, sheathing-leaves which arise from the peri-
phery of the axis, and these, as well as all the subse-
quently formed leaves, derive their vessels from the axis,
the very earliest development of vessels must be peri-
pheral ; and this method of succession is preserved, as
long as any leaves continue to be formed. Moreover,
the upper end of the vascular bundle, says the author,
tends towards the base of the leaf, the lower end becomes
elongated obliquely downwards in the form of a delicate
filament, consisting of parenchyma, but which never
extends into the root. The spots from which the vas-
cular bundle arise at the summit of the bud, are organi-
cally predetermined : they are there placed with their
upper ends converging obliquely towards the centre, and
become elongated in both directions i. e. grow both
downwards and upwards. The spot at which the upper
end of the vascular bundle passes to the leaf, is situated
either on the same side of the stem as that in which the
course of the vascular bundle is chiefly included, or oppo-
site the point of origin of the vascular bundle, obliquely
as regards the diameter ; in which latter case, the vas-
cular bundle traverses the entire stem obliquely. As the
summit increases in length and thickness, each vascular
bundle crosses another bundle, either in the interior of
the stem or nearer to the periphery, at the point where,
either ascending almost perpendicularly, or suddenly taking
a horizontal direction outwards, it enters into the leaf.
These are, undoubtedly, the most valuable remarks which
have been made upon this subject, and I am glad to
find that they confirm what I have stated previously,
though less completely. Still I must confess that I am
in doubt regarding the growth of the vascular bundles
upwards and downwards. The author's explanation does
not contain any proof of the occurrence of this growth
in both directions. In my opinion it always occurs
upwards, but in the same manner as is seen in the corm
at the base of the young stem, except that the bundles
cross at smaller angles as the growth of the stem pro-

PHYSIOLOGICAL BOTANY. 255

ceeds. Some of the bundles may occasionally diverge
more considerably, as the author and Mirbel agree in
stating. I have no doubt that here also new vascular
bundles are formed between the old ones, which certainly
takes place as in Dicotyledonous trees.

On Gaudichaud's Theory of the Merithals. By Pro-
fessor GUIS. MENEGHINI. Giornale Encyclop. Italiana,
vol. i, p. 17. This memoir, which was written as early
as 1843, at the time of the meeting of the Congress of
Philosophers at Lucca, must have contributed much to
draw attention to Gaudichaud's system. The author
sketched the elements of this system, which consists in
the unity of the axial system of the plant with the appen-
dicular system, the plant being regarded as composed of
phytons, intermediate forms, so to speak, between the
stem and leaf. The author admits this system as esta-
blished, endeavours to illustrate it by analogy with
animals, and considers that it must have the greatest
influence upon organography. It would have been more
desirable that the author, with the acuteness he possesses,
had investigated the whole theory more closely. He
would then have seen that the exposition of the system
rests upon an arbitrary assumption, which can only pro-
duce hypothetical results. That the radicle of Dicotyle-
donous seeds, of the Grasses and Cyperoidese, is the future
stem, has been known for thirty years ; but this is not
the case in other Monocotyledons. That all parts of an
organic being are originally one, no one can doubt, but
that the latter is directly developed into these parts, and
that the developed leaves, e. g., do not constitute the
entire stem, is evident on the slightest examination. The
author states his explanation of Gaudichaud's system to
be as follows : The fibres neither ascend nor descend ;
they are formed in the previously-existing cellular tissue,
by a gradual conversion of the cells of the parenchyma ;
the organization of the fibres is determined by the cur-
rents of the nutritive fluid and the descending fluids (sap) ;

256 PHYSIOLOGICAL BOTANY.

the mechanical action of these, and the materials which
they bring with them, contributing to it, &c. But this
conversion has not been proved to occur ; in all proba-
bility it is entirely false, and the currents of a fluid may
become suddenly changed; in the germinating cotyledons
of the Monocotyledons, when emitting radicles, they pass
suddenly upwards into the stem, and downwards into the
root. But it is perhaps unfair to criticise the author from
an old memoir, as since that time he has proceeded with
his investigations, and we have still much to expect
from him.

New Researches upon the Development of the Axis and
Appendages of Plants. By M. C. NATJDIN. Annal. d.
Scienc. naturell., 3 ser. vol. i, p. 162. These observa-
tions are in general accurate and valuable, although they
are not new. The leafy parts (appendages), says the
author, are the lateral products of an axis, which at first
consist of cells only ; they also at first consist of cells
only, not containing vessels, and the apex of the axis, the
centre of a bud, exhibits a tubercle (mammelon) which is
in connexion with the pith. The second part of my
Select Anatomico-botanical Plates contains many figures
which show this more distinctly than the author has done ;
but so it is, our labours run parallel with those of fo-
reigners, yet we are generally a little before them. But
no; the author is actually acquainted with Duchartre, Guil-
lard, and Schleiden, who have investigated this subject.
Recently, in the second part of my ' Anatomy of Plants/
the plates contain further illustrations of this subject.
What he states of a few Monocotyledons only, viz. that
the spots, where the vessels arise, are denoted by a mo-
dification of the cellular tissue, applies to most plants,
and has also been illustrated in the part of my work
alluded to above. The distinction which the author
makes between the axis and the foliaceous parts, viz.
that the former grows at the extreme point, whilst no
addition takes place at the extremities of the latter, is

PHYSIOLOGICAL BOTANY. 257

not perfectly correct, for no addition is in fact made to
the very extremity, or external circumference of the axial
parts, any more than to the apex and the upper end of
the borders of the leaves. The author confounds this
with another subject, namely, that the leaf appears and is
developed before the petiole ; and in support of this view
he quotes Morren, who (in opposition to me) has asserted
that this does not occur in the aquatic plants, e. g.
Hydrocharis morsus ranee ; but when the entire plant is
examined, the commencement of the leaves is distinctly
seen before any trace of the petiole can be distinguished.
The author's remarks upon the development of Monoco-
tyledons are very imperfect. He only treats of the bulb
of Narcissus pseudo -narcissus, and that in a very super-
ficial manner. He might also have seen very strikingly,
in the first part of my ' Select Anat.-botan. Plates,' that
the vascular bundles are continued from the stem into
the root. His idea of distinguishing a spadix from a
spike is to the purpose ; thus, in most cases, the summit
of the bud is covered by leaves, whilst there it grows up
naked.

Micrometric Researches upon the Development of the
Elementary Parts of the Annual Stems of Dicotyledons.
By M. G. HARTING. Ann. d. Sc. Nat., 3 ser. vol. iv,
p. 210. It is difficult to give an extract of this extensive
and valuable memoir, without exceeding the limits of a
Report like the present. The author only treats of the
annual shoots of Dicotyledonous plants. First, of the
method and means adopted in his micrometric investiga-
tions. Then of the opinion that an annual shoot may be
regarded as composed of several individual internodes
(merithals) of different ages, but having the same original
structure, so that, by the examination of the various
internodes of the same shoot, we may conclude as to the
changes undergone by any internode in the course of its
growth. The youngest internode of the shoot, it is well
known, is the last, and even a superficial examination

258 PHYSIOLOGICAL BOTANY.

shows, that the lower internodes first cease to grow.
Investigations upon the growth of the shoot of Tilia
parviflora follow, arranged in tables ; these include : in-
crease of the individual internodes in length ; increase of
an individual internode at different ages ; increase of the
pith ; multiplication of the cells of the pith ; increase of
the longitudinal and transverse diameter; of vascular
and liber layers ; transverse diameter in proportion to
the longitudinal; parenchymatous layer of the bark;
number of rows of cells, proportion of the diameter of
this layer to the diameter of the internode, increase of
the cells of this layer in comparison with the increase of
the cells of the pith ; Schleiden's callenchymatous layer,*
i. e. the layer of remarkably long cells, which exists
beneath the epidermis in many plants ; number of cells
in the peripheral layers. We next have similar investiga-
tions upon Humulus Lupulus, as also upon the nucleus
(cytoblast) in the cells of the pith, the corpuscles existing
in the layers of the liber, and the growth of a shoot,
which had been deprived of its leaves at the apex. Again,
investigations upon the shoots of Aristolochia Sipho, Phy-
tolacca decandra, and Sempervirum arborescens. Then
follow the results : 1. The growth of each internode de-
pends upon the formation of new cells, the expansion of
the cells, and the thickening of their walls. 2. The multi-
plication of the cells takes place in three directions : in that
of the radius radial growth ; in that of the periphery
peripheral growth ; and in that of the axis longitudinal
growth. 3. The radial multiplication only occurs in the
buds. 4. This multiplication is produced by means of cross
septa, which are formed in the previously existing cells,
without the latter becoming subsequently absorbed ; the
subdivisions thus formed continually become more and
more isolated, in consequence of their expanding on all
sides. 5. The expansion of the cells in the radial direc-
tion is uniform and equal, so that the diameter always

* " A term as elegant as it is superfluous," as Schleiden once said against
me on a similar occasion.

PHYSIOLOGICAL BOTANY. 259

retains the same proportion until the formation of wood
occurs. 6. The lignifying layers (vascular layers and liber)
do not begin to expand in a radial direction until the
fibrous cells have begun to acquire thickness ; but then
with a force which exceeds that with which the pith and
the parenchyma of the bark expands. 7. During this
period, the cavities of the cells and vessels expand uni-
formly, and this also still takes place after the thickening
of the fibrous cells has commenced. The increased space
which the vascular and prosenchymatous layers occupy in
the old segments, must be ascribed to this increase in
thickness, not therefore to deposition upon the inside of the
wall. The author says in a note, that the development of the
fruit in the Drupacese, and of the albumen in the seeds
of some Monocotyledons, shows, that the increased thick-
ness also arises from deposition upon the outside of the wall.
Must the expansion necessarily arise then from a thickening
of the walls ? The interstitial growth of cells and vessels,
which undoubtedly takes place in old stems, indicates an
expansion without any thickening. 8. The lateral expan-
sion of the cells composing the different layers, usually
takes place (at least in the pith, in the parenchyma of the
bark and of the epidermis) with the same force in all
directions. But this law is liable to exceptions according
to the activity of growth. 9. In the stems of those
plants in which no central canal is developed (Tilia and
Aristolochia) , the cells constituting the pith, the liber,
and the parenchyma of the bark, do not multiply peri-
pherally, but only in the direction of the longitudinal
axis. The peripheral multiplication is only perceived in
the epidermal and callenchymatous layers. 10. In the
plants just mentioned, neither the number of the vascular
bundles, nor, consequently, that of the vessels increases.
The diameter of the latter increases in proportion to the
expansion of the vascular layers. 11. In those plants, on
the contrary, in which a central canal is developed, the
cells of all the layers multiply peripherally, as is also the
case with the vessels. This multiplication, in the author's

260 PHYSIOLOGICAL BOTANY.

opinion, causes the absorption of the fluids in the internal
cells, and their desiccation, which explains the origin of
the inner cavity. 12. When canals containing gum occur
in the pith or in the parenchyma of the bark (Tilia), they
increase but little in diameter during their growth, but
they multiply, and when the elongation is completed, they
again diminish and become thickened. They already
exist at the very earliest period. 13. In those stems in
which a central canal is not formed, the increase in breadth
depends upon the radial expansion of the cells, excepting
in the layers of the callenchyma and the pith. In those
stems in which a central canal occurs, the part which
the multiplication and expansion of the cells takes in its
formation, is variable. 14. This is also the case in the
longitudinal growth. 15. The longitudinal multiplication
of the cells, as also their expansion, takes place simulta-
neously at all points of the internode, but in those inter-
nodes which are still elongating, the cells of the pith, of
the parenchyma, of the bark, and of the epidermis at the
apex of the internode, are shorter than those at its base,
and the latter again are shorter than those at the apex of
the adjacent older internode. When the expansion of
the cells at the base has ceased, that of the cells at the
apex still proceeds for some time. 16. The smallest cells
multiply most, hence the epidermal cells more than those
of the parenchyma of the bark, and the latter more than
those of the pith, but no definite proportion is observable.
17. Whilst the internode is still very young, the growth
takes place principally by the multiplication of the cells
only. When the internodes of a plant, after they have
acquired their full length, are of slightly different lengths
(Tilia, Humulus, and Aristolochia) , the numbers of the
cells of the pith and bark, in the younger internodes, form
a geometrical progression. Moreover, the younger the
internodes are, the less they grow, and when the growth
is accelerated with advancing age, it observes a geome-
trical progression. All this shows that the multiplication
of cells takes place in geometrical progression. For

PHYSIOLOGICAL BOTANY. 26 I

instance, each cell subdivides into two, and each of these
again into two more, and so on. As the internodes be-
come older, the growing process becomes still stronger,
because the expansion of the cells is then combined with
their increase in number. The growth decreases gradually
at last, because after the increase in the number of cells
formed has ceased, the expansion still continues to some
extent. 18. Hence three principal periods may be distin-
guished in the growth of the annual shoots of Dicotyle-
dons : 1st, that in which the intern ode still constitutes a
part of the bud, and where the increase in the number of
cells is only radial ; 2dly, that in which the internodes
increase both in length and breadth, and this again, a t
arising from the increase in the number of cells only, or
b y from simultaneous increase in the number and the
expansion of the cells, or lastly c, from the expansion of
the latter alone ; 3dly, that in which the growth in the
axial direction has ceased, but where the lateral expansion
still continues. 19. Since the longitudinal diameter of
the cells in those internodes in which the elongation has
ceased remains the same, the various different lengths of
the internodes must arise solely from the formation of a
larger number of horizontal layers. The author ascribes
the differences which are perceptible, when the effect of
season upon growth is considered, to this circumstance,
sometimes more, sometimes feAver of these layers being
developed. 20. In the cells of the pith and of the paren-
chyma of the bark of the youngest internodes, in which
the growth occurs slowly from the multiplication of cells,
we find a substance consisting of very small globules.
Very few of the cells are furnished with a nucleus (cyto-
blast) containing a corpuscle. On the other hand, we
find in many cells small groups, or simply rings, consist-
ing of these globules. On examining the adjacent older
internode, we recognise in a large number of cells, and in
the next internode (in which increase in the number and
expansion of the cells are simultaneously taking place) in
all the cells, perfectly-developed nuclei, which are quite

262 PHYSIOLOGICAL BOTANY.

transparent and provided with their corpuscles. In the
transverse section, they appear to be situated in the centre
of the cells, whilst in the longitudinal section they are
generally seen to be adherent to the wall. They have a
flattened form, hence they are detected with difficulty in
this aspect. At this period, the granular matter has for
the most part disappeared. In the youngest internodes,
which have ceased to elongate, and commonly in the next
internode also, nuclei still exist in a few of the cells, but
they are usually situated upon the lateral walls of the cells.
They disappear in the older internodes. 21. During the
earliest period of the growth of the stem, neither the pro-
duction of new cells, nor their expansion, nor the thicken-
ing of their walls is dependent upon the presence of the
terminal bud, or upon the leaves existing at the end of
the articulation.

These excellent investigations might form the basis of
the theory of the growth of plants. It is very desirable
that similar investigations should be made upon Dicoty-
ledons, the sterns of which are subdivided by nodes, and
then upon Monocotyledons. The merismatic increase in
the number of cells in those plants which the author has
examined appears to me proved. There can be no doubt,
however, that the septa which are produced must be
double ; the manner in which they are formed still re-
mains to be determined. In many cases, the growth
certainly does not take place according to a geometrical
progression ; in these an interstitial growth of cells must
occur, perhaps in combination with merismatic division.

It is completely decided by all the investigations which
have been made, that no formation of cells within cells
ever occurs either during the growth, in length or thick-
ness, of vegetable structures, unless we regard merismatic
division as such, which would be incorrect. It is by no
means my intention to deny the occurrence of this mode
of formation in those cases where perfectly new bodies or
parts are formed, and the formation of the young plant in
the embryo-sac furnishes an instance of the production of
cells within cells.

PHYSIOLOGICAL BOTANY. 263

The curvature of stems towards the light will be re-
ported upon in an article in which the general action of
light upon plants will be discussed. Dutrochet has made
some observations upon the movements of the free points
of plants furnished with tendrils, which were spoken of in
the previous Annual Report. In the Compt. rend., 1844,
ii, 295, there are some observations, by the same author,
upon the Movements of the free Points of twining Plants.
They take place in the same direction as that in which
the stem coils itself. Dutrochet connects them with the
spiral position of the leaves. In Solanum Dulcamara the
winding of the stem is sometimes from right to left, some-
times the reverse, and the spiral of the leaves is double.
I shall here merely observe on this point, that Mohl has
already observed the curvature of both the stem and
the tendrils when unsupported. Dutrochet also gives
the times in which the turns are completed ; but they do
not appear to be very constant.

Dutrochet has also made the observation, that in Epilo
bium molle Lam., (E. parviflorum Schreb.) some of the stems
grow directly into the earth, like roots. They are thicker
than the upright stems, and are furnished with more
cortical substance, which Dutrochet ascribes to their de-
scent into the earth, or rather the thickness of the cortical
substance arises from the moisture of the earth. (See
Compt. rend., 1845, ii, 1186.)

Boucherie reports, that sections of wood prepared by his
method (see Annual Report for 1840, pp. 360, 384), kept
sound in the earth for three years, whilst other unpre-
pared sections of the same wood entirely rotted in the
same place. Compt. rend._, 1845, ii, 1153.

As regards WYDLER'S Morphological Communications,
Bot. Zeit., 1844, 641, 657, 688, 705, I shall merely
remark, that they do not permit of condensation into the
form of an abstract. The author there assumes, as his
basis, an ingenious method of explanation, which was
proposed by Al. Braun (Flora, 1842, 694). But other

264 PHYSIOLOGICAL BOTANY.

expressions should be selected, instead of such indistinct
and incorrect ones as uni- and bin -axial, because these
are very obscure. In these investigations Wydler has
especially endeavoured to explain the remarkable struc-
ture of the Solanacese. It is so marked that the Natural
Order is recognised by it, however the same occurs in
other Natural Orders and individual genera, for instance,
in the Boraginaceae, Phytolacca, and others.

ROOT. TUBERS. PRICKLES. TENDRILS. GLANDS. STOMATA.

On the Tendency of Roots to strike into the Earth (but
really into mercury only). By PAYER. Compt. rend.,
1844, i, 993. In the year 1829, says the author, Pinot
remarked that some seeds of Latliyrm odoratus, which
he had caused to germinate upon mercury, forced their
radicles into it. This penetration was afterwards con-
sidered to arise merely from the weight of the seed;
others did not observe any penetration ; and De Candolle
regarded the penetration as arising from the stiffness of
the root. Payer then made some experiments upon the
point, and found that the radicles of Polygonum Fago-
pyrum, although they are stiff and thick enough, remained
on the surface, whilst, on the other hand, the much more
delicate roots of Lepidium sativum penetrated to a con-
siderable depth. Nor does the weight at all contribute
to the effect. If the root be withdrawn from the mercury,
it does not again penetrate the metal, but it sometimes
shoots further, and then the new portion does so. Light
and heat increase its power of penetration. The author
thinks that the power possessed by the roots of penetrating
the earth arises from their power of avoiding light, and
seeking for proper soil. This is explaining something
that is unknown by something else which is still more
unknown. We have only given an extract from the
memoir.

The results, only, of a memoir, by Durand, upon the

PHYSIOLOGICAL BOTANY. .265

same subject, occur in the Compt. rend., 1845, i, 861.
He first historically adduces Pinot's observation, and
adds, that Dutrochet ascribes the phenomenon merely to
the pressure of the seed ; he then speaks of Mulder's in-
vestigations, which were made at the same time, and
which prove the contrary. He next details the results of
his own experiments. When the seeds are fixed above
the surface of the mercury, the roots penetrate ; but if
this is not done, they do so only when the seeds are
placed at the side, between the glass and the mercury, or
when a layer of the organic matter is deposited from the
water, which fixes the young plant. The seeds of Poly-
gonum Fagopyrum do not yield any of this matter to the
water, hence the roots do not penetrate.

The Report of the Commission upon these two memoirs
is given in the Compt. rend., 1845, i, 1257. Several
points which in the memoirs are not described, but merely
alluded to, are more detailed. The reporter, Dutrochet,
finds fault with Payer's memoir, because he has not
stated whether he fixed the seeds above the mercury, and
if so, how this was done. The following remarks are
quoted from Durand's memoir : When the seeds of
Poly gonum Fagopyrum are properly fixed during germi-
nation, the radicles always penetrate the mercury. If
the seeds are placed in water over mercury, without being
fixed, they lose as much in weight as the water which
they displace weighs ; hence they exert less pressure upon
the mercury, and therefore cannot penetrate it. If, under
these circumstances, they are but slightly covered, they
penetrate to a certain extent. As has been already stated,
the semi-solid layer of precipitated organic matters fixes
the little plant at the surface of the mercury, and replaces
the artificial fixation. As the seeds of buck- wheat do not
yield any organic matters to the water, we need only add
a little of some extract to the water to produce the same
result. Some experiments performed by the reporter
(Dutrochet) follow next. He says, "we have used several
kinds of seeds in these experiments, and especially those

266 PHYSIOLOGICAL BOTANY.

of Latliyrus odoratm ; but we have never seen that the
radicles of these seeds sank more deeply into the mercury
than was caused by the pressure which the weight of the
seeds exerted upon the radicles, i. e. not more than three
millimeters." The Report concludes with the statement
that the phenomenon ensues according to known laws ;
that M. Durand has discovered that the penetration of the
radicles into the mercury depends upon the seeds being
fixed, and that when this is not the case, the seeds merely
penetrate as deeply as the pressure of the seed causes
them to do. The Report censures M. Payer for inaccu-
racy in the description of his experiments ; but this would
apply still more strongly to the account which the reporter
gives of his own experiments, for he makes no mention
of the direction of the radicles which penetrated, and yet
this is the main point, if it were really the pressure of the
seeds which forced the radicle into the mercury. More-
over it will appear easily explicable that the radicles
penetrate the mercury, when the seed is fixed, because
this is the only remarkable circumstance, and it is strange
enough, when Durand's statement is brought forward.
If seeds are placed in water over mercury, without being
fixed, they lose as much in weight as the water which
they displace; thus they exert less pressure upon the
mercury, and cannot, therefore, penetrate ; since, by
fixing them, the weight is entirely removed, and could
not, therefore, cause the roots to penetrate, there is no-
thing else but the force of the onward growth to make
the root descend, and it is remarkable that this is not
prevented by mercury. The experiment of Payer is very
remarkable ; he found that the roots of Latliyris odoralus
descended through several layers of mercury., in an in-
genious apparatus arranged for the purpose. It is also
remarkable, that when the radicle is withdrawn from the
mercury, the portion which had penetrated will not again
do so ; but the newly-emitted part does, an experiment
which excludes all mechanical explanation. The experi-
ments upon the penetration of the radicles of unattached

PHYSIOLOGICAL BOTANY. 267

seeds germinating upon mercury, do not appear to me of
any importance.

A remark which has already been frequently made, is
repeated by H. Jaubert, in the Compt. rend., 1845, ii,
360, namely, that on that side of a tree on which the
branches are strongest, strong roots also exist. He says
that he has very frequently found this to be the case in
digging up trees in Sologne. It may be well to recall
to mind, since these observations correspond with the
view, that the nutritive fluids ascend through the spiral
and dotted vessels, and as these vessels do not anastomose,
they ascend from the root in a straight direction. That
the branches, however, assume curves, like the roots, an
instance of which is here afforded, appears to be acci-
dental.

TREVIRANUS has described a remarkable Formation of
Tubers, in Sedum amplexicaule. D. C. Bot. Zeit., 1845,
p. 265. In this plant, says he, the new shoots which
are destined for reproduction are much thickened for
about the length of an inch at the summit ; the leaves at
this part are also closely crowded, whilst at the lower
part of the shoot they are scattered. About the period
of the summer solstice, not only the main stem, which
has flowered, dies, but also the lateral shoots, the thick-
ened points of which constitute the above newly-formed
living shoots. On examining the latter, we find, com-
pletely incased by the dried sheath-like basis of the
leaves, a cylindrical mass of cellular tissue, the cells of
which contain granules of starch, and the centre of which
is occupied by a small ring of fibres and vessels ; at its
point is a bud consisting of the rudiments of a few leaves,
and which is marked with the scars of fallen leaves. It
is a tuber, formed by the confluence of closely- crowded
leaves. About the middle of August these tubers emit
new leaves ; these clothe the next year's stem, which ter-
minates in a flower; they are not, however, sheathing
like those which surround the tubers, but semi-cylindrical,
like the leaves of Sednm acre, reflexwn, &c.

268 PHYSIOLOGICAL BOTANY.

PIETRO SAVI on the Prickles of Amaranthus spinosus.
Giorn. Encicl. Ann., 1, t. i, pp. 17, 310. The author
considers that these prickles are not stipules, as has been
thought, but that they are the lowermost, early-developed
leaves of an axillary branch. The author's view is per-
fectly correct ; they are situated in the axil of the leaf,
low down on the axillary branch, and the main proof
consists in their producing tufts of flowers in their axils,
which is never the case with stipules. It would be very
remarkable were stipules to occur in one species of
Amaranthus when they are not found in any other, nor in
genera related to it.

On the Tendrils of the CucurbitacecB. By ATTILIO
TASSI. Giorn. Encicl. A. 1, t. i, P. 2, p. 382. In op-
position to the view that they are stipules. His view is
principally founded upon the instance of Sicyos Buderoa
Hook., the alternate leaves of which are furnished on one
side below the base with either one, three or six filaments,
three or four only of which in the latter case acquire com-
plete development. The author also alludes to the remarks
made upon them at the Italian Scientific Congresses.
Auguste St. Hilaire (Memoir, d. Musee, t. ix, p. 192),
whom the author does not mention, is the authority for
their being considered as stipules. He quotes an instance
of Ulaterium, and a case of abnormal development of
Cucurbita Pepo, which had produced stipules instead of
tendrils. On this point I have remarked, in my * Ele-
ments of Philosophical Botany/ vol. i, pp. 318-9, "But
the so-called stipules in the Gourd were furnished at the
point with a small tendril ; hence the tendril (as is fre-
quently the case with spines) had produced leaves.
This minute tendril appears to be absorbed in Elaterium ;
for the true stipules never arise from one side only of a
leaf; they are very rarely stalked, and the stalk when
present is never round, as the tendril is almost invariably.
The tendril, which is now the subject in question, is also
situated close to the branch like the spine, and is also a
supernumerary branch."

PHYSIOLOGICAL BOTANY. 269

A. St. Hilaire also treats of this point in his ' Morpho-
logic Vegetale/ pp. 185-6, and says, in opposition to the
view that the tendril occurs on one side of the leaf, that
we find on one side of the leaf a developed stipule, and
on the other an abortive one (Ervum monanthos), and that
it is but a small step from this to its complete absence (?).
Moreover, in one of the Cucurbitacese in the garden at
Paris, he had noticed two tendrils. My ' El. Ph. Bot.'
were published as early as 1837, the 'Morphologic' in
1841. M. Tassi must extend his inquiries beyond what
is done in Italy.

New Researches upon the Structure of Cistomes. By
GUGL. GASPARRINI. Naples, 1844, 4. The author has
previously described a pouch or sac, which is adherent to
the stomata internally. He calls these sacs cistomi, be-
cause they are attached to the stomata (stomi). In the
present short communication he now describes a canal
which arises from the sacs. His investigations were
principally made upon Cactus Peruvianus, then upon
Ornitliogatum nutans, and Arum Italicum. I have also
examined Cactus Peruvianus, and have seen the sacs but
not the canal, which the author himself has only figured
in some, not in all. But the epidermis must be strongly
boiled with hydrochloric acid to exhibit the sac ; hence it
appears to be nothing more than the internal membrane
of the aerial reservoir, which the thicker epidermis (cuticle)
has entered and lined, as has already been remarked by
Mohl. Mohl has also found that the epidermis sometimes
extends into the cellular tissue, and there forms, as it
were, canals. Too strong treatment with acids materially
destroys the connexion of the parts, so that the true
structure is no longer recognisable ; and this is the case
in the present instance. I have not been able to find the
canal which the author figures from Ornithogalum nutans.

C. MULLER has communicated to the 'Bot. Zeit./
1845, p. 793, Some observations on the Resinous Exudations

270 PHYSIOLOGICAL BOTANY.

of the Birches. Beneath the epidermis a small heap of
cells, filled with green matter, is seen, and which is
slightly elevated above the surface ; it gradually becomes
larger, and lacerates the epidermis. More cells then
become superimposed upon each other, and form a capi-
tule, with a more or less thick pedicel formed by the
lower cells. The outer cells subsequently become wholly
converted into a resinous matter, and are surrounded
with a dense brittle mass, the pedicel still remaining
unaltered. Finally, the granules escape from the epider-
mis. The dense brittle mass dissolves in alcohol or ether,
forming a mucous mass, leaving no trace of any membrane
(which however might very easily be concealed in the
mucous mass). The author gives the chemical exami-
nation of betuline ; he considers it to be a kind of
stearoptine.

LEAVES.

Remarks upon the Arrangement of the Leaves in Dicoty-
ledons. By K. S. KUNTH. Bericht d. Akad. d. Wiss. z.
Berlin, October 1843. The position of the leaves coin-
cides with that of the buds, says the author, and when a
bud is about to be formed, a portion of the pith is forced
through the woody substance towards the surface of
the stem. The spot where this takes place depends upon
the arrangement of the woody bundles; thus the first
year's shoots of the oak are pentagonal, and the leaves are
also arranged in five ranks. On endeavouring to connect
the leaves by a line in the shortest course, this can only be
done in a spiral direction, and from left to right ; more-
over the spiral line, to be enabled to reach the nearest
leaf, must pass over one of the angles of the wood, to
arrive at a leaf which belongs to the same series. Five
angles are not always present, still five divisions of the
wood may always be assumed to exist when the leaves
are arranged in this manner. The author next refers the
arrangement of the leaves in the shoots of Castanea vesca

PHYSIOLOGICAL BOTANY. 271

in two ranks to the five-ranked, remarking that, when we
commence with the posterior single odd rank of leaves,
the fourth and third are developed, whilst the first, second,
and fifth are not formed. In the same manner he re-
duces the three-ranked arrangement of the leaves of Alnus
glutinosa to that in five. From the alternate (scattered,
says the author, which is, however, the opposite of
fascicled} leaves he comes to the opposite leaves, which he
regards like the former, not, as at the same height, lying
in a transverse section made perpendicular to the axis,
but merely approximated and alternating. He proceeds
in the same manner with the verticillate, or whorled
leaves. This valuable contribution to the theory of the
arrangement of leaves deserves great attention ; and it is
certainly of importance to consider the angles of the stem
in connexion with the arrangement of the leaves. We
must connect with this

On the Arrangement of the Parts of Flowers. By
K. S. KUNTH. In the Bericht d. Akad. d.Wissen. Berlin,
Feb. 1844. The whole of the elements of a perfect
flower, says the author, form several depressed, equally-
divided whorls, and may be connected either by one or
two spiral lines which run parallel. Hence we must dis-
tinguish uni- and bi-spiral flowers. The organic spirals
of Dicotyledonous flowers consist typically of five-mem-
bered, bi-spiral whorls ; but there are uni-spiral flowers.
These usually consist of three whorls ; the sepals form
the first whorl, the stamens the second, and the pistil the
third. These are the only true apetalous flowers, the
other apetalous flowers which occur being readily distin-
guishable by the number and position of the stamens ;
to these the Thymeleaceae, Polygonacese, &c. belong. The
flowers of Monocotyledons are distinguished from those
of the bi-spiral Dicotyledons, merely by the three-mem-
bered whorl, and thus, like the latter, they also present
a calyx and a corolla ; hence it is incorrect to ascribe to
them a perigone. I shall merely remark here, that this

272 PHYSIOLOGICAL BOTANY.

expression emanates from Ehrhart, and signifies the pre-
sence of both calyx and a corolla. The word is well con-
structed. P. externum signifies the calyx, P. internum
the corolla ; hence the expression may be conveniently
used where an intermediate form is present, as in many
of the Monocotyledons, especially, however, in the Thy-
meleacese, the Polygonacese, &c., for the true calyx of a
Chenopodium is very different in structure from the calyx
of the corolla of a Daphne.

Disquisition upon a Problem propounded in Phyllotaxy.
By ANTON. PHESTRANDREA. Messina, 1843. A. M.Argen-
tano presented for solution, in a journal (Interprete Ann.,
iv, No. 7), a problem in the theory of the arrangement of
leaves, and it is very satisfactory to find that this German
theory has travelled as far as Sicily ; which would cer-
tainly not have been the case, if the excellent report of
Martius and Bravais had not appeared in the ' Annales
des Sciences Naturelles.' The problem is as follows :
In a plant, the arrangement of the leaves of which is
spiral, the spiral winds 13 times around the stem, and
the angle of divergence amounts to 137^ degrees ; to find
the number of leaves or foliar organs which compose the
cycle. The solution is very simple. If we denote the
angle of divergence by d, the number of turns by a, the
number of foliar organs in the cycle by m, we have, ac-
cording to Schimper, d = , in which we may

denote one of the three magnitudes an unknown oc. We

Qn* 1 ^

then have 137+ 11= ou , whence (137 + $ x =

360-13, and therefore x= 34. The object is to bring
Schimper's theory under consideration (although Al. Braun
only is mentioned), in which the divergence of the gene-
rating spiral is assumed as 137^ degrees, according to
Bravais. The author comments at some length upon the
aid which one science can afford another ; as an instance

PHYSIOLOGICAL BOTANY. 273

of which this is adduced, and the example here calculated
out for beginners.

Schimper's representation of the Arrangement of
Leaves is undoubtedly very ingenious, because it collects
the vague expressions of the spiral arrangement of leaves
into a comprehensive review. The formula given above
must be regarded as the fundamental formula, from
which the others may be deduced. Its application to
opposite and whorled leaves, the leaves of axillary bf anches,
even the development of the leaves in the buds, and the
parts of flowers, is no less ingenious. Schimper's ex-
planation is somewhat awkward, and it was therefore very
important that Al. Braun detailed this system more accu-
rately, more copiously, and more clearly. An excellent
memoir next appeared by MM. L. and A. Bravais, in the
'Ann. des Scienc. Natur./ 2d ser., t. vii, pp. 42-110.
The authors examined the spiral positions of the leaves
and foliar parts, the secondary spiral lines, as they are
represented on the developed surface of a primary cylinder,
where, namely, the spiral lines running from right to left,
and those from left to right intersect each other; and
point out as the foundation of the whole theory, that
when the numbers of the above two rows of spiral lines
are respectively primary numbers, there exists a spiral
line, which includes all the spots at which the leaves are
attached, a generating or including spiral, but if they have
a common divisor, verticillate positions occur. In the
first case, the angles both of the separate spirals (se-
condary spirals) and of the individual members in the
spirals will coincide with the horizontal line, and the
secondary divergences with the divergence of the pro-
ducing or general spiral. If we call the number of a
member in a secondary spiral line n, the divergence of
this spiral dn, the divergence of the general spiral d\ 9
and m the number of turns made by this spiral before it
arrives at n, we then have ndl = m . 360 + dn. This
formula serves for the calculation of the divergence of the
general spiral. It is then found, by direct observations,

274 PHYSIOLOGICAL BOTANY.

that this divergence in most cases amounts to 137 30' 28",
an irrational angle ; in some other and more rare cases,
the angle, which is also irrational, is = 99 30' 6", or
77 57' 19", or 151 8' 8". None of these angles are
altered, at least their mean values, either by the dissimi-
larity of the members which follow each other or other
local circumstances. The increase is worthy of note,
at least according to its average value, especially in
Schimper's method of finding the angle of divergence,
since we cannot always meet with a foliar part occurring
exactly in a vertical line above it. It is remarked also,
that for this purpose, the outer rind often requires to be
removed in order to distinguish the false from the true
angles. The authors also extend their observations to
the false whorls, they show that the including generating
spiral extends to the underground stem, that the direction
of the spiral is the same in the stem and branches, but
exerts no influence upon the direction of twining stems.
The convergence of two spirals into one, which is some-
times noticed, may arise from the abortion of one spiral
or a confluence of two spirals into one, an entire series
may then be absent, whence the existence of several series
becomes doubtful. It appeared to me important to refer
to this memoir again, since it appears to be read less than
it deserves, for it not only contains a great many theo-
retical considerations, but also numerous investigations
made upon the plants themselves. Meyen's remarks upon
this subject, which have been given in previous Annual
Reports, do not appear to me altogether to the point.

In my 'Elem. Philosophise Botanicse,' pp. 450-1, I
endeavoured to discover a general expression for the ex-
positions given by Schimper and Braun, by which they
might be more easily reviewed. I was unacquainted
with the memoir of Bravais ; it appeared in 1837, simul-
taneously with my ' Elemental I started from Schimper's
theory. Let m represent the number of leaves (including
in this term bracts also) between two leaves which come
next each other in a vertical line, but which are arranged

PHYSIOLOGICAL BOTANY. 275

around the stem in a turns. If we project them in a
circle, the distance between two leaves which are nearest

one another, is equal to an angle of , the apex of which

is in the axis of the stem ; but if this circle be drawn out

a times, the angle becomes This is Schimper's law,

according to which, a spiral including all the leaves is
taken, and the circumference of the circle is made = 1 .
The line which converges towards or is parallel with the axis
of the stem, between two leaves situated in this line, we
shall call the principal line, because it is that with which
we set out in the investigation. Then to determine the
position of any leaf or member in the entire generating
spiral, we must ascertain its distance from the principal
line. The first member, as we have just shown, is situated

at the angle , the second at , the third at , and so
m m m

on, which, deducting each angle from 360 or 1, gives

the series 1 -, 1 , 1 , &c. Thus altogether
m m m

m a m 2a m 3a m na m ma .,,

m m m m m

which the series terminates, because there are only m
members present. As in this determination of the dis-
tance, the entire circumference of the circle is passed
through several times, we must leave out these circuits in
calculating the numbers, to find the true or least distance.
Let m = 21, = 8, as Al. Braun found for the cone of
the Fir, these numbers, disregarding the signs become

13.5.3.10.2.6. 7.1.9. 4
4.9 1. 7.6.2.10,3.5.13

Thus the numbers recur in the second half, as follows from
the form of the series, and when m is an odd number,
the mean number is doubled. Where m=5, a = 2,
which is most common, we have 3.1.1.3, whence it is

276 PHYSIOLOGICAL BOTANY.

at once evident that the angles are twice passed over, if
the stem be at all regularly pentagonal and the leaves
situated at the angles. When m loses itself in na, the
series is at an end even before any of the leaves or mem-
bers have become included in the spiral, because the

quotient then becomes a multiple (in even numbers) of ,

to the fundamental primary angle of the distance of
one leaf from another, whence one leaf lies in a straight
line above the other, and forms with it the principal line.
Twenty-one leaves may be placed in 2. 4. 5. 8. 10. 11. 13
turns of the generating spiral, but not in 3. 6. 7. 9. 12,
because these numbers might yield a product na, in which
m = 21 loses itself, namely 7.3.7.6.3.7.9.7.2.7.
This is not the place for pointing out the application to
the secondary spirals, the number and properties of which
may be easily deduced from the fundamental series, as
has been done in the work quoted above. It appears to
me that this series most easily allows of our comprehend-
ing all cases of the arrangement of leaves, and I have
therefore again brought it forward, and have given some
parts fully and distinctly.

Naumann has had his Memoir in Poggendorf's ' An-
nalen/ upon the Quincunx as the fundamental law of the
arrangement of Leaves, printed separately, merely with
the correction of the errors of the press. It was spoken
of in the last Annual Report.

The Polarity of Buds and Leaves. By MAX. WICHURA.
Flora, 1844, p. 161. Perhaps the author's meaning
may be best comprehended from the following pas-
sages : " If, from the point where one bud is situated,
we wish to arrive at the place of the next above it, this
may always be effected in two different paths. One
ascends in a direction to the right, the other to the left.
If this be tried with any stem, the buds of which are
separated from each other by the divergence J, it is a
matter of perfect indifference whether we take the right

PHYSIOLOGICAL BOTANY. 277

or the left path, for both are of equal length. But
whenever the divergence is different from this, one path
must be shorter than the other. The next question,
then, is 1. Do the buds upon this stem, when they are
imagined to be all connected together, either on the longer
or the shorter path, follow in the same direction above
one another, so that the connecting line presents one
continuous spiral, or is this not the case? 2. Which of
the two paths unequal in length runs towards the right,
and which toward the left ? The decision of the former
of these questions teaches us, that in addition to
numerous plants, in regard to which it must be answered
in the affirmative, there are some in which the direction
of the spiral is reversed at every point where a bud is
attached. Therefore, when we there denominated the
connecting line a continuous spiral, we shall here, from
analogy with what in geometry is called a fractional
degree, apply to it the name of fractional spiral.
Examples of this arrangement are afforded by the biserial
buds, as in part of the Papilionacece, in Tilia, Celtis,
Cercis, Ulmm, Carpinus, Corylus, Morus, Statice, Be-
gonia, Phyllanthus, and many others. I have before me
a branch of Tilia grandifolia, and find an arrangement
of the leaves, not at all uncommon, namely, ; and ac-
cording to the fundamental series, the divergences of the
individual members are 2 1 4 * 7 ' 10, thus one small
and three great distances, whence the leaves appear almost
biserial ; but they are by no means really so, for they
distinctly lie in a continuous spiral. The author pro-
ceeds : " The system of the continuous spiral is, there-
fore, altogether distinguished from that of the fractional
spiral, not only by the direction in which the buds follow
one another, but also by the inner structure of the buds
themselves. Buds which are developed in the same
direction one above the other, surround the stem on two
or more sides, are placed under each other in an uniform
relation, which frequently degenerates into irregularity.
This is the condition of the indifference buds. But those

278 PHYSIOLOGICAL BOTANY.

which succeed each other in a continually alternating
direction in two series, separated by less than half the
circumference of the stem, are symmetrical, and the pro-
duct of forces uniform, but acting in opposite directions,
and this is the state of polarity. But all the buds and
leaves upon the stem grow upwards, and I cannot see how
polarity can act here. It always acts in directly opposite
directions, and not at angles. Important as it is to con-
sider an object not individually and separately, but as a
whole, there is no polarity manifested in this case, unless
we change the meaning of the word. The primary phe-
nomenon is ascent in the direction of a spiral, from a ver-
ticillate position.

Morphological Communications, by WYDLER; on the
Characterization of Foliaceous Structures external to the
Flower. Bot. Zeit., 1844, p. 625. Wydler here treats
of some of Schimper's definitions of leaves. He divides
the leaves on a plant into inferior leaves (Niederblatter) ,
frondose leaves (LaubUatter), and superior leaves (Hock-
blatter) ; again, each leaf into a vaginal portion, petiole,
and lamina. The frondose leaves consist of a, vaginal
leaves, consisting of the vaginal portions only, as in the
Iris ; b, petiolar leaves, consisting of a petiole only, as in
the Acacias, Indigofera juncea, and Lathyrus Aphaca ;
c, frondose leaves, consisting of vaginal portion and
petiole, as in Allium Cepa ; d, frondose leaves, consisting
of petiole and lamina, as in most plants ; e, laminar
leaves, consisting of the lamina only, as in sessile leaves ;
/, frondose leaves, consisting of vaginal portion, petiole,
and lamina, as in Arum, Palms, Rheum, the Urnbelliferse,
Leguminosae, and Rosacese. Many of these details are
applicable. The name vagina,, or sheath, is not inappro-
priate ; and by means of it, we can easily express the
distinction between an entire and a half sheath. Instead
of the word lamina, which is not German, we have plate,
leaf-plate (blade). The leaves of Iris do not consist
merely of a vagina, but of vagina and leaf-plate. Allium

PHYSIOLOGICAL BOTANY. 279

Cepa is also furnished with a leaf-plate, as is seen in
young leaves. The subdivision, <?, shows that the whole
section gives but an indefinite and unpractical summary,
for the relations of the principal nerves and their distri-
bution is entirely unnoticed.*

In this Gazette, for 1844, i. 134, there are some
Observations upon the Growth of the Organs of Vegeta-
tion, considered in a Systematic Point of View, by A.
GRISEBACH. They were made upon Phlox paniculata,
Dianthus plumarius, Saxifraya hypnoides, Peucedanum
Alsaticum, MenyantJies trifoliata, Aristolochia Sipho, and
Ampelopsis hederacea. To these are added some remarks
upon the growth of stipules. As the observations, so to
speak, are special, they cannot be reduced into an ab-
stract. There is a supplement at p. 345, which treats of
the vegetating points in the sheaths of the leaves. Sin-
gularly enough, parent-cells and secondary cells within
them are here spoken of, although, at p. 138, the author,
from his observations upon Phlox paniculata, arrives at
the conclusion that in it the longitudinal growth of the
lamina is effected by Mohl's cell -division.

Upon the Occurrence of Sugar upon Leaves. By Pro-
fessor VON SCHLECHTENDAL. Bot. Zeit., 1844, p. 6.
The author especially describes the glands secreting sugar
in Viburnum Tinus ; they exist at the margin of the leaves
near the base, one on each side, projecting like a short
tooth. When the plant is kept in a room in the winter,
a white lump of sugar arises on the surface of these glands.
Since the lump of sugar in Viburnum Tinus, as also in
Rhododendron Ponticum and Clerodendron fragrans, are
only observed in plants living in a room, the author
supposes that the saccharine liquid is solidified by the
desiccation.

* The reporter gives but an imperfect idea of this paper. The Nieder-
bldtter or inferior leaves are the bud-scales and analogous structures, the
frondose leaves are the general leaves of the stem, the Hochbldtter or supe-
rior leaves, the bracts, &c. Eugl. Ed.

280 PHYSIOLOGICAL BOTANY.

On the Saccharine Glands of Leaves. By UNGER.
Flora, 1844, p. 703. In many Acacias, as A. longifolia,
armata, verticittata, and myrtifolia, the author saw a
saccharine fluid drop away ; and on careful examination,
found at the base of the phyllode, close to the thickening
at its upper margin, a small punctiform depression, which
is the excretory duct of a slit-like cavity in the substance
of the phyllode. This cavity is surrounded by peculiar
cells with thin walls, which collectively form a glandular
apparatus, in which the saccharine fluid collects, and
from which it is gradually evacuated. There are two
vascular bundles connected with the saccharine glands,
and giving branches to them, the vessels of which are in
short joints and curved, and in this manner are lost in
the parenchyma of the circumference. The author sub-
joins some observations, principally relating to the honey-
like secretions from the leaves and branches caused by
insects.

On the Propagation of Cardamine pratensis L., by
Means of its Leaves. By JUL. MUNTER. Bot. Zeit.,
1845, p. 537. The author accurately describes the de-
velopment of young plants from the leaves of Cardamine
pratensis, for the most part after Cassini, whose accuracy
Schleiden has rendered doubtful. The hemispherical
nodule, from which the plants are developed, occurs at
the spot where the three principal nerves of the leaflets
radiate from each other into the leaf. The roots spring out
on the upper side, at first grow upwards, and afterwards,
when they have acquired sufficient length, descend. In
addition to this, a second bud frequently arises from the
middle of the central rib. The most remarkable point is
the confirmation of Cassini's observation, that the leaves
of Cardamine pratensis separate, live under water, and
there emit young plants. The author found that the
chlorophyll disappeared then, and considers justly that it
probably serves for the nutrition of the plant.

M. Pietro Savi has also observed the development of
young plants upon the leaves of Cardamine pratensis in

PHYSIOLOGICAL BOTANY. 281

the garden at Pisa, and briefly describes them in a note
to Meneghini and Savi's ' Memoir upon the Appendages
of the Leaflets of Acacia Cornigera/ in the ' Giorn.
Enciclop.' i, p. 406. These appendages occur upon the
point of the leaflets, but only of the lower ones ; they are
absent in the upper ones towards the point. They are
elliptic-elongate, of one sixth or one eighth part of the
length of the leaflet, of a whitish-yellow colour, and are
furnished with a central nerve, which is a prolongation of
the nerve of the leaf. Around the central nerves spiral
vessels occur ; the rest consists entirely of cellular tissue.
He then treats of the morphological character of this
appendage, and of the view that they are probably abor-
tive buds, according to Gaudichaud's theory, in which
the leaf is considered as a phyton. Although it cannot
be denied that the leaves may be considered as such,
when Cardamine pratensis is adduced as an example,
they must rather be regarded as degenerations of the
extremity of the teeth of the leaves themselves, or their
entire margin (come degenerazioni deW eslremita delle
dentellature delle foglie stesse e del loro totals). Were
this not the case, they must be considered as glands,
which view, however, is opposed by their constant posi-
tion at the margin of the leaves, and especially at their
most prominent points. The appendages are evidently
so-called glands, which do not secrete any fluid. Their
morphological character, in my opinion, is an indication of
a tendency of the leaf to become again pinnate. Perhaps
the authors entertain the same opinion.

KIRSCHLEGER has described the stipules of Platanus.
Flora, 1844, p. 725. These structures, which have been
long known, are merely described by the author, because
Endlicher says, when speaking of the PlataneaB, stipulce
mdlce. But Endlicher is right ; they are not stipules, but
ochrece, such as exist in the Polygonese, &c. They are
not situated at the sides of the petiole, but surround the
axis above the base of the petiole.

282 PHYSIOLOGICAL BOTANY.

FLOWERS FRUCTIFICATION.

Contributions to our Knowledge of the Inflorescence of
Cannabis, Humulus, Urtica, and Parietaria, as also ofPar-
nassia palustris, Erodium, and Inipatiens. By WYDLER.
Flora, 1844, p. 735, 757, 759. They contain accurate
explanations, and form supplements to the memoir in the
'Linnsea/ 1843, which was spoken of in the last annual
Report ; also remarks upon the ramification of the latter
plant. The author's observations upon the arrangement
of the leaves of Polycarpon tetrapliyllum, should be com-
pared with the above. Flora, 1845, p. 33.

Remarks upon the Symmetry of the Corolla. By D.
WYDLER; Bot. Zeit, 1844, 609. The author's mor-
phological investigations are communicated in a very
indistinct style. " As we know," says the author, " most
symmetrical corollas may be divided by a line into two
equal halves, which, starting from the derivative axis of
the flower, we imagine to be drawn through the middle of
the upper odd sepal and the lower odd petal towards the
bract. To this class, amongst others, belong Pinguicula,
Utricularia, the Labiatse, &c." But the corolla is a solid
figure, which cannot be divided into two equal halves by
a line, although it can by a plane. The author means to
say that a tranverse section of the flower near its base, a
plan of the flower, is divided by a line into two equal halves.
This is the manner in which the subject is illustrated
by the figures. With a different mode of illustration,
numerous conclusions might be drawn, but these cannot
be given here.

Morphological Considerations upon Arduina bispinosa.
By PIETRO SAVI. Giornal. Encicl., i, 113.

Remarks upon some Microscopic and Superficial Organs
of Plants. By PIETRO SAVI. Giornal. bot. Italiano,

PHYSIOLOGICAL BOTANY. 283

i, 27. The author describes the papillae and their con-
tents, which exist upon the flowers of Chrysanthemum
indicum, Thunb. ; he considers them as glands. I do
not find it mentioned that these papillae have long since
been found and described upon ail true corollas. The
blue powder upon the leaves of Chenopodium and Atriplex,
is incorrectly referred to them ; it consists of globules
of wax.

On merismatic Formation of Cells in the Development
of Pollen. By Dr. F. UNGER; 1844. An excellent
memoir in a few pages. " According to my observations,"
says the author, " the earliest traces of renewed organiza-
tion in the mature parent-cells, appear as very thin and
delicate streaks, which either run transversely across the
centre, or laterally, according to the position of the parent-
cell. These streaks, as any one may easily satisfy himself
by turning round the parent-cell, are nothing else than
extremely thin and transparent walls, which divided the
uniform granular matter into several parts. These walls,
which must necessarily be formed from the above-men-
tioned contents, are so perishable, that they dissolve in
water, which renders it probable that they consist of gum.
But simultaneously with this phenomenon, a spontaneous
separation of the granular mucilage occurs, which espe-
cially tends to show that from this moment the nucleus of a
cell begins to be developed in each portion. The forma-
tion of these walls still proceeds, so that they not only
soon acquire greater firmness, but also greater thickness.
The first commencement of true membranous development
(for the earlier deposit can scarcely be considered such)
distinctly takes place from the walls towards the central
point. First, there appear projecting ridges, and from
these the membranes crystallize, as it were, more and
more toward the interior, so that we can trace the progress
step by step." Further on : " Hence there are no special
parent-cells, separate and inclosed by the parent-cells, but
only special parent-cells which originate as divisions of

284 PHYSIOLOGICAL BOTANY.

the parent-cell, and only undergo partial separation in
the highest stage of their development/' Hence the
result is that, even in pollen, the formation of cells from
a cell-nucleus never occurs.

In the 'Flora/ 1844, p. 359, FACCHINI communicated
the Investigations of AMICI of Florence, upon the impreg-
nation of the Embryo which are opposed to Schleiden' s
theory of the development of the embryo. Schleiden did
not omit at once answering them, 1. c. 787. Eacchini
therefore gave the Italian text of Amici s memoir, as it
exists in the Transactions of the ' Scienzati' of Padua, with
the remark, that all present were convinced by Amici.
Schleiden, 1. c., 593, then accuses all who were present
of gross ignorance, and abuses Aimer's figures in his
peculiar style. Any one who wishes to know how Amici,
the discoverer of the pollen-tube, is treated, may read this
paper. We gladly turn from this subject to a remarkable
work

Experiments and Observations upon the Organs of
Fructification of the more Perfect Plants. By C. FR.
GARTNER, Stuttgard, 1844-8. We have here such an
abundance of experiments and observations made with
great calmness and circumspection, that we may assert
with truth, that no modern work has contributed so
much to the physiology of plants as the present. This
is not the place for carefully going through the whole, we
can only enter generally upon it, and notice a few of the
various points on which it treats. Moreover, in addition
to the new and peculiar results which it contains, we have
in every case a notice of the opinions of others, which are
contradicted or confirmed by reasoning and experiment.
1 . On the Flower. Cause of the abortion and falling off of
the flower. 2. Of the Calyx. If the impregnation of the
ovary has not been effected, the calyx decays, and assumes
a diseased appearance ; it remains in this state for several
days, according to the species of the plant. 3. On the

PHYSIOLOGICAL BOTANY.

285

Corolla. Castration has no effect upon the corolla, and
the presence of the stamens is not in the least necessary
for the preservation of its integrity and perfect develop-
ment. The styles are generally developed subsequently
to the corolla, in a few plants only does the reverse occur,
as in Lychnis diurna, vespertina, Dianthus barbatus and
superbus. When in the latter case the stigmas are sprinkled
with the pollen of the same plant, whilst the flower is but
little or only half developed, the growth of the latter is
checked, or entirely ceases. Many observations and ex-
periments are made upon the diurnal sleep of plants;
impregnation exerts great influence upon it. 4. Upon
the secretion of honey (nectar) in flower. Tending rather
to contradict the views which have been asserted, than to
propose definite laws. 5. Upon the stamens of plants.
The observation is remarkable, that hybridation gives a
tendency to wasting of the anthers. The duration of
the power of the pollen is very different in different plants,
and also very different from the duration of the suscepti-
bility of the female organ to impregnation. The author's
remarks upon the pollen-tubes, and their penetration into
the micropyle, do not bear the least relation to the rest.
6. On the evolution of heat from flowers. Many original
observations. It occurs also in the female organs, and
is often connected with the odour. 7. On the pistil.
8. On the phenomena of irritability and motion in the
flowers and organs of fructification of plants. A number
of observations and experiments, especially upon the irri-
tability of the pistil in Mimulus. Development of the same.
When cut off and kept in moist sand, they exhibited
the same phenomena as when in situ, the destruction of
the one stigma exerts no particular influence, concussion
produces no effect upon them. Experiments with che-
mical irritants. Among these there are also experiments
with oil of morphia (a mixture of morphia and poppy- oil),
which prove, that the irritability and capability of move-
ment of the pistil of Mimulus is diminished and finally
destroyed by it. This is also the case with Strychnine

286 PHYSIOLOGICAL BOTANY.

oil. Castration has no further influence upon the irri-
tability, than that it lengthens the period of duration of
the flower, and thus of the pistil also. Experiment upon
the action of the pollen of the same flower ; this is only
exerted during the time of the susceptibility to impreg-
nation, but chemical irritants act at other times. In
many flowers, movement occurs during the period of im-
pregnation, without irritability; observations upon the
deportment of the flowers of Tropaolum majus, &c.
Observations upon Stylidium. 9. On the impregnation of
perfect plants. The dehiscence of the anthers in many
plants occurs uniformly before the flowers open, but in
most, afterwards. Action of light, heat, and moisture.
The author never succeeded in obtaining ripe seeds from
branches of dicotyledonous plants which were cut off and
kept in water. Other promoters of impregnation. Forty
grains of pollen were required to produce impregnation in
Malva Mauritiana. Similar experiments upon Tropceo-
lum majus. Precautionary rules and phenomena of arti-
ficial impregnation. The nucleus is capable of continuing
its growth for some time without impregnation, but it
does not produce an embryo. Phenomena observed after'
fructification in the nucleus and the seeds of twelve
flowers of Lychnis vespertina. Similar observations upon
Staphylcsa pinnata, during a period of four months ; in
both cases with accurate anatomical investigations, but
without figures. He arranges my observations upon
Angraecum with his own upon the embryo of Corydalis,
but he is still unacquainted with the figures in my
Anatomico-Botanical Plates, which would have taught him
the great difference between them. 10. Upon the abor-
tion of flowers, fruits, and seeds. Shorter than the other
memoirs. 11. Upon the production of fruits, with seeds
capable of germination, without the application of pollen.
The observations of others upon the subject are criticised,
and their insufficiency pointed out. His own observations
yielded a perfectly negative result. 12. On the impreg-
nating power of plants. 13. On false impregnation.

PHYSIOLOGICAL BOTANY. 287

The author follows Kolreuter in applying this term
(afterbefruchtung) to abortive impregnation, produced
by the pollen of the same plant. 14. On the effect of
sprinkling the stigma with foreign matters. Experiments
of the author in opposition to Henschel's now forgotten
experiments. We anxiously look forward for the second
part.

A Paper, by ROPER, on the Flora of Mecklenburg, 2 pt.
Rostock ; contains Researches upon the Flowers of the
Grasses, which we recommend all botanists to read. It
is especially opposed to Schleiden's theory, viz., that the
lower and outer valve of the glume, or the palea inferior,
forms a trifoliar perianth with the upper and inner valve
of the glume, which originally consists of two valves.
It distinctly shows how Schleiden's censoriousness mis-
leads him into the greatest inconsistencies. It also con-
tains several other investigations of importance. As on
most points I agree with the author, it would be super-
fluous to make any remarks. Nor is this the place for
explaining the true condition of the flower of Lolium temu-
lentum (Crap alia, Schrank), regarding which the author
appears to be mistaken. One observation more. The
author sets philology at defiance, and adopts such ex-
pressions as sepalum, tepalum, &c. Language is so re-
markable and wonderful a production of the human mind,
that it must not be trifled with; an unfortunate proceeding,
which has in recent times been especially revived by
De Candolle.

'On the Signification of the Lower Glume of the Flower
of Grasses. By HUGO VON MOHL, Bot. Zeit., 1845,
p. 33. The author also shows by an analysis of the
common monstrosity found in Poa Alpina, that the lower
floral glume is not to be regarded as a perigonial leaf, but
as a bract.

Note upon the Organography of the Flower oftheMal-
vacece. By M. DUCHARTRE. Compt. rend., 1844, i,

288 PHYSIOLOGICAL BOTANY.

p. 487; 1845, i, p. 349. Report, ibid, ii, p. 417, and
in detail, in the Ann. de Scienc. Natur., 3 ser., t. iii,
p. 123. Report, p. 150. The external calyx (involucre),
is first formed, then the true calyx, from a single piece.
A globule next arises in this, which exhibits five tubercles;
this soon subdivides into two parts, and thus the flower,
even in its youngest condition, is furnished with ten
staminal tubercles. Five small folds next appear close to
the calyx; these are at a tolerable distance apart, and
constitute the commencement of the petals. Hence the
flower is at first pentapetalous. The development of the
stamens, internally, next follows, according to a double
plan, first by concentric circles, which grow inwards, and
then by the deduplication of the stamens. They are really
situated opposite the petals, but in many Malvaceae, we
find that the filaments become elongated above the sta-
mens, and form five teeth, which alternate with the five
groups of the andrsecium, and thus represent the inner
row of stamens. The formation of the pistil varies, and
the author admits four different methods by which it
takes place. To this subject belong :

Observations upon the Organogeny of the Flower, and of
the Ovary in particular, of Plants having a free central Pla-
centa. By M. A. DUCHARTRE. Comptes rendus, 1844,
i, p. 1105. Development of the flowers of the Primu-
laceae. The calyx is first formed in one piece, not several, as
Schleiden states. Five tubercles next appear, from which
the stamens are formed; the appearance of the corolla seems
to precede that of the stamens, when they alternate with
the segments of the corolla; otherwise, it follows after. The
pistil appears simultaneously with the corolla in the form
of a cone, and the placental tubercle fills up the ovary.
The ovary next rises up and forms the style. The summit
of the placenta subsequently begins to elongate, and passes
into the canal of the style, and is not, therefore, at first
connected with the stigma. The report upon this me-
moir, by Ad. Brogniart, Ach. Richard, and Gaudichaud,
is, on the whole, favorable.

PHYSIOLOGICAL BOTANY. 289

The Observations of GELEZNOFF upon the Development
of the Flower of Tradescantia Viryinica. Bullet, de la
Societe Imper. des Naturalistes a Moscou, vol. xvi, 1843;
Flora, 1844^). 144 ; Bot. Zeit., 1844, p. 183, should be
compared TOn the above.

The Academy of Naples has given an account of the
Memoirs which it has received in reply to its Programme
upon Caprification. The Memoir No. 1 denies its in-
fluence in fertilization. Female flowers are always found
in the fruit, but no males; and the impregnation
of the figs remains a mystery. The author does not
recommend caprification. The Memoir No. 3 arrives at
the following conclusions : 1 . The wild fig is not the male
of the cultivated fig, as it has been considered. 2. Inas-
much as the structure of the flower and the seeds in the
varieties of the cultivated fig are exactly the same, there
appears no reason why caprification should be requisite in
some varieties and not in others. 3. The insect does not
hasten the ripening, neither does it contribute to the
setting of the fruit any more than it does to its impreg-
nation. 4. The falling off of the fruit of the wild fig,
which contains no larvae, proves nothing, for when many
fruits have set upon the tree, they still fall off, even when
larvae are present. 5. The cause of the falling off must
be sought in other circumstances ; in the climate, changes
of the weather, &c. 6. Caprification is perfectly useless,
either for ripening or setting the fruits. The Memoir
No. 5 contains the conclusion : That the action of the
Cynips upon the cultivated fig is entirely mechanical, and
merely serves, like any other irritant, to accelerate some-
what the ripening of the fruit. Hence, when this is not
requisite, caprification is perfectly useless, nay, even in-
jurious to the perfect maturation of the fruit. The
Memoir No. 6 considers caprification requisite, but only
in the case of the abortive figs. One memoir only, to
which, however, we shall only briefly allude, considers
that it is necessary for fructification. In my early days

290 PHYSIOLOGICAL BOTANY.

I had an opportunity of observing caprification in Portugal,
and in the account of my travels I have stated that capri-
fication exerts no influence upon impregnation. How-
ever, many varieties become larger and more beautiful
when they are pierced by this minute CynifSJi, as is very
truly stated in the Memoir No. 5.

In the 'Thuringian Horticultural Gazette/ Nos.l and 2,
Prof. Bernhardi treats of Bastard Forms. He now con-
siders that the so-called bastard forms of the genus
Gymnogramwa (Ceropteris] might arise, not from impreg-
nation, but from the coalescence of the roots with each
other, because they germinate in hot-houses in numbers
together. As an instance, he mentions a plant of Cytisus
Adami, which was produced by grafting C. purpureus
upon C. alpinus, whereby a hybrid was produced, which
frequently assumed the characters of a bastard, and often
returned to its primitive conditions, at one time producing
purple, at another yellow flowers. This is remarkable
enough, and is the first instance of the formation of
bastards in this manner.

FRUIT. SEEDS. GERMINATION.

Memoir upon the Development and Characters of true

Ji -L /

and false Arils. By J. E. PLANCHON. Montpellier, 1844.
An excellent contribution to our knowledge of the
changes undergone by the seeds in their young state.
First, a history of the meaning of the word aril. Then
an investigation of the nucleus in Passiftora. Since, in
this instance, an expansion of the umbilical cord is not
formed until after impregnation, as it is only connected
with the seed by means of the external umbilical aperture
(hilum), and at the opposite end is widely open, since,
therefore, this structure agrees with the generally admitted
notion of the word aril, the author denominates it a true
aril. The aril of Euonymus latifolius is very different,
although it agrees with the former in many particulars.

PHYSIOLOGICAL BOTANY. 291

After the fall of the petals and stamens, the nucleus
grows somewhat : a protuberance is then formed at the
margin of the exostome, which grows out, expands into a
membranous margin, and extending towards the base of
the nucleus, forms a hemispherical cover, which covers the
base of the nucleus, but leaves the micropyle completely
uncovered, whilst, on the other hand, the true aril covers
the micropyle. The author calls the aril of Euonymus a
false aril or arillode. The definitions then of these parts
are : the true aril is an accessory covering of the nucleus,
which is developed around the umbilical aperture (hilum),
in the same manner as the proper coverings, and either
covers the exostome, or would do so if it were sufficiently
developed. The false aril or arillode is an expansion of
the margins of the exostome, which is reflected around
this aperture, but always leaves it uncovered. We have
examples of true arils in the Dilleniacese, the Samydaceae,
the Bixinese, Nymphcea ccerulea and alba, but it is absent
in Nuphar lutea. Moreover, Chamissoa is mentioned as
an example, and a description is then given of the seed
of Cytinus Hypocistis. The ovary of this plant is filled
with a mucus, and upon its walls there are ramified but
compactly superimposed placentae. I shall give the descrip-
tion of the ovule and seeds in his own words : " Ovula
ortholropa, creberrima, minutissima, occidua, utrinque at-
tenuata, basi arillata. Integ. unicum, vasculis destitutum,
arete adhcerens, membranaceum, pellucidum, apice perfora-
tum. Nucleus solidus, cellulosus, ovulo conformis, subdia-
phanus. Arillus irregulariter cupuliformis, brevis, crassus,
margins incequalis e cellulis laxis latis constans, vix quar-
tam ovuli partem inferiorem oUegens, ab eodem facillime
secedens. Semina (in fructu siccato) ovulis conformia,
pallide lutea, mucilagine in lacrymas solidas, vitreas co-
agulata involuta. Arillus et integumentum ut in ovulo,
prior won raro obliterates. Nucleus solidus, omnino eel-
lulosus. Embryo nullus." The author in fact thinks, that
no embryo is present, because as the ovule is orthotropous,
impregnation could only occur through the mucus of the

292 PHYSIOLOGICAL BOTANY.

ovary. But may not the entire nucleus be an embryo ?
He includes amongst false arils the remarkable envelope
of the seed in Opuntia, the formation of which is here
shown to occur from two lateral expansions of the funiculus.
Moreover, the false aril of Euonymus latifolius, which has
been already mentioned, belongs here ; the tubercle in
the Euphorbiacese also is only the thickened margin of
the exostome, and the aril, as it is called, of the Polyga-
lacese agrees in many points with it. In Clusia flava we
must assume, that the outer envelope of the nucleus,
which is simple in the greater part of its extent, becomes
doubled, divided into two unequal prolongations beyond
the exostome. It is doubtful whether the Nutmeg does
not belong here. The author denominates by the term
Strophiola the glandular excrescences situated along the
raphe ; they are independent of the umbilical cord and
the exostome, and he mentions, as an example, the seeds
of Arum Canadense. Finally, the history of the nucleus
of some species of Veronica, especially V. liederapfolia and
V. Cymbaria, with remarks upon the genus Avicennia.
The peculiar operculum of the seed of the latter originates
from the embryo-sac, which in Veronica is converted into
albumen. The embryo- sac of Avicennia ruptures the
nucleus in the ovary, and the embryo also ruptures the
embryo-sac by too rapid germination in the fruit. In
Veronica hedercefolia, the nucleus at an early age is
reduced to the embryo-sac only, and has no covering.
The details must be obtained by the examination of the
author's work itself.

In a memoir which M.Guglielmi Gasparrini read before
the Academy of Naples as early as ] 842, he endeavoured
to show, that the fruit of the Opuntia is merely a branch
adapted for this purpose. The nuclei are at first situated
in the central cavity in rows, the wall of the cavity is
not a distinct organ, like the ovary in other plants, but
consists of a special complicated fibrous tissue, which is
produced for this structure. This fibrous tissue is both

PHYSIOLOGICAL BOTANY.

293

podospermous and trophospermous. The free podosperm,
although very short, is the primary membrane of the
ovule ; after impregnation they become covered with
cells (otricelli), which arise from the growth of the sur-
rounding cellular tissue, and constitute the pulp, by
means of which the seeds become separated from one
another, and are lost in the pulp. The pulpy mass con-
taining the seeds is not connected to the receptacle or to
the summit of the flowering branch, but to the upper
part of the bark, where the petals, stamens, and the outer
styles arise, by means of a fibrous tissue which descends
so as to terminate in the seeds. This subject deserves
more minute examination, not only in regard to the
adhesion of the style to the ovary, but especially with
regard to Planchon's investigations, which appear to form
a supplement to those of Gasparrini.

Note upon the Embryogeny of Taxm baccata. By MM.
DE MIRBEL and SPACH. Compt. rend., 1844, i, p. 114.
In addition to the embryo which is developed, the authors
found two vesicles ; they do not consider these as abortiv*
embryos, for long before the embryo appears, these vesicles
become adherent by their base to the apex of the embryo-
sac, and the tube (boyau), which is above each of them,
becomes elongated by means of the nucleus almost as far
as the surface of the upper end. Hence the authors con-
sider that they serve in impregnation.

Investigations upon some Vegetable Monstrosities, which
may serve to illustrate the origin of the Pistil and of the
Nucleus. By AD. BROGNIART. Compt. rend., 1844, i,
p. 513. It is a question whether the seeds arise from
the axis, or from the margins of the carpellary leaves.
' The example which it is my intention to make known,"
says the author, " exhibits in its carpels all the stages of
the formation of the leaf ; it exhibits at its margin ovules,
some of which scarcely differ from the normal ovule;
others form imperceptible transitions to lateral lobes of
the carpellary leaf. If is taken from a plant of Delphi-

294 PHYSIOLOGICAL BOTANY.

nium elatum, which flowered in 1841 in the Garden at
Paris.

On the Development of the Ovary, the Embryo, and the
anomalous Corollas of the Ranunculacece. By BARNEOUD.
Compt. rend., 1845, ii, 352. As the rows of stamens
become doubled, we find at their base two oval, some-
what approximated plates, which alternate with the sepals;
and a little more internally, in a different plane, we find
five other ovate plates, which are smaller than the former,
and opposite the segments of the calyx. This shows that
the two spur-shaped petals belong to another and larger
whorl, the other elements of which abort regularly ; the
next whorl also aborts. The ovule is always anatropous,
but exhibits three remarkable types. According to the
first, it makes half a rotation upon itself, in an horizontal
direction, and the exostome is directed towards that side
on which the placenta is situated, as in the Helleboreae,
Pceonieae transverse anatropy. In the second type, the

Jr / / XT y

ovule rotates vertically, and the margin of the exostome
is turned towards the base of the carpel inferior ana-
tropy, as in the Ranunculese. According to the third,
the ovule is suspended, and the exostome turned towards
the summit of the cavity superior anatropy, as in the
Clematideae and Anemoneae. The embryo-sac exists pre-
.viously to impregnation ; it becomes filled with cells,
which are subsequently transformed into the albumen.

Chemical Researches upon the Ripening of Fruit.
Compt. rend., 1844, 784. This subject is a most im-
portant one, and as the chemical changes which occur
during the ripening of fruits are very distinct, perhaps one
not very difficult of attainment. But isolated remarks
against this and that point, such as we find here, are of
no use. The experiments must first be performed upon
one fruit only, and cherries would be best for this pur-
pose, because they quickly ripen, and undergo great
changes during maturation ; moreover, as it appears to
me, their analysis would be easier than that of the pear, &c.

PHYSIOLOGICAL BOTANY.

295

Germination of Charophyllum Bulbosum. By Professor
KIRSCHLEGER. Flora, 1845, 401. The seeds had ger-
minated in spring, with two cotyledons ; however, no
bud existed between the cotyledons ; but at the base of
the plumule a tuber was developed, which, during the
same year, bore radical leaves, and in the following
year flowers and fruit. This circumstance was not un-
known, having been long since observed in Bunium bulbo-
castanum.

An account of some Seeds buried in a Sandpit, which
germinated. By WILLIAM KEMP. Annals of Nat. Hist,
v, 13, p. 89. The layer of sand which contained the seeds
was nearly 25 feet (see Ann. Nat. Hist.) below the sur-
face. They germinated, and were found to be those of
Polygonum Convolvulus, and a variety of Atriplex patula,
with Eumex acetosella, an Atriplex, &c., only common
British plants. The author considers them to be of an
immense age, and supposes that the Tweed had flowed
through the valley, and deposited the seeds, before a large
vein of trap-rock passed through it. Probably a more
accurate examination would somewhat diminish the time.

Wahlberg's remarks in the Report of the Swedish
Academy are more reasonable. (Vide Mora, 1845, p. 61.)
He sowed the seeds of several plants, foreign and Swedish.
The place was covered with building materials for many
years, and when these were removed and the soil dug up,
several plants sprung up which had previously flowered
there.

Periods of flowering and ripening of several wild
and cultivated plants, , which were observed in 1843, for
the purpose of forming a scale of the development of
plants in various parts of the duchy of Nassau, in the
Annual Reports of the Society for the promotion of
Natural Science in the Duchy of Nassau, by Dr. K.
THOM^E. Wiesbaden, 1844. The plants were: Eibes
ruhrum and Grossularia, Fragaria vesca, Rosa canina,
Primula veris et officinalis, Sambucus nigra, Primus

296 PHYSIOLOGICAL BOTANY.

spinosa, domestica, avium, Pyrus mains, Secale cereale,
Triticum vulgare, Hordeum vulgar e, Avena sativa, Solatium
tuberosum, Vitis vim/era, Juglans regia, and Castanea
vesca.

INDIVIDUAL ORDERS AND GENERA OF THE PHANEROGAMIA
IN REFERENCE TO PHYSIOLOGY.

Description of the Female Flower and Fruit of Eafflesia
Arnoldi, ivith Remarks on its Affinities, and an Illustration
of the Structure of Hydnora Africana. By R. BROWN.
Transact, of the Linn. Soc., vol. xix, pt. 3 (1844) p. 221.
The author, with his usual accuracy and well-known
acuteness, investigated the above objects, and treats of
them with a certain heartiness which renders the subject
very attractive. The whole is illustrated by the excellent
drawings of Ferdinand Bauer. The ovarium of Hydnora
may be regarded as composed of three confluent pistilla,
having placentae really parietal, but only produced at the
top of the cavity. It would, however, certainly be diffi-
cult to reduce Eafflesia to this type. The author then
describes the development of the ovules of Eafflesia in
their earliest state, which agrees with that occurring in
Phaneragamous plants generally, the lower portion of the
papilla becoming dilated, forming a cup, and enveloping
the future integument and the nucleus. Thus the author's
description differs, and rightly, from that of Mirbel's.
According to the author, a curvature occurs, as in several
Phanerogamia, but only in the upper part of the funiculus,
whereas in the Phanerogamia generally, the curvature is
produced in that portion of the funiculus, which is con-
nate with the testa. The reason of this may be, says
R. Brown, that the testa is absent in the seeds of
Eafflesia. The author only found pollen-tubes in Cytinus.
The testa of the seed of Eafflesia is evidently the same
which exists in the unimpregnated ovule, and is very
hard ; the inner membrane is thin ; the nucleus ap-
pears to be entirely composed of cellular tissue, but in

PHYSIOLOGICAL BOTANY.

297

the middle of it a cylindrical portion is found, which con-
sists of large transparent cells, which the author regards
as the embryo. The seed of Hydnora resembles that
of Eafflesia in many points. Its nucleus consists of a
dense albumen, in which a spherical embryo is found.
In Cytinus the seeds are minute, and retain at their base
the bipartite membrane, which may with most probability
be considered as a prolongation, of the testa. The latter
is easily separable from the nucleus, which appears to
consist of an uniform cellular mass, as in the Orchidacea3.
He finally asserts that vascular tissue is not absent in the
Rafflesia and Balanophorese, and adds, that frequently,
plants which are very different in external aspect, are of
the same internal structure ; I might add, as in Cycadese
and Coniferse. Lastly, he gives a botanical description
of the female flowers and fruit of Eafflesia Arnoldi and
Hydnora Africana.

On Macrozamia Preissii. By G. HEINZEL. Nov.
Act. Acad. Leop., vol. xxi, pt. i, p. 203. This forms an
inaugural dissertation, which appeared at Breslau, and
well deserves to be received into this work. The de-
scription of the plant is very good, and great attention is
paid to the physiology. The stem and leaves are not
treated of, although these are of great importance, but
only the male and female organs of generation. It is a
very remarkable circumstance, that the hard unilocular
anthers are irregularly produced from the scale in most
species. The author describes very accurately a minute
pedicle upon which the anther is formed ; I have detected
it in other Cycadeae, and always find it to contain a vas-
cular bundle. The author then advances an adventurous
morphology, according to which the scales merely con-
sist of convoluted stamens (filaments). These strained
hypotheses should not be had recourse to in matters
relating to the vegetable kingdom, and no morphology at
all is far better than a forced one. With regard to the
ovule, he does not follow R. Brown, but thinks that what

298 PHYSIOLOGICAL BOTANY.

he considers to be an exostome is rather a stigma. A
detailed Criticism of this paper, by Dr. Gottsche, of
Altona, is given in the Bot. Zeitung., 1845, pp. 366,
377, 398, 413, 433, 447, and 507, which contains much
remarkable matter, and therefore deserves great attention.
Thus it contains a minute examination of the ovule of
Encephalartos lonyi/olius, with comparative observations
upon other Cycadea3 and Coniferse. We cannot follow
the author in his investigations, for this would require a
separate memoir.

Upon the Structure of a full-grown Stem of Cycas cir-
cinalis. By F. A. W. MIQUEL. Linnaea, 1844, p. 125,
tab. 4, v, 6. A good description of a full-grown living
stem, of which we were not before in possession. The
internal structure is especially remarkable ; it consists of
a cortical parenchyma, which is composed of three layers
of cells. The wood is divided into concentric, unequal
and irregular layers, which are separated from each other
by thicker or thinner layers of parenchymatous cells, con-
taining starch. Each woody layer is divided into almost
quadrangular or club-shaped woody portions by distinct
medullary rays. On examining the large woody layers,
we find that they take a very serpentine course. Those
vessels which assume a lateral direction, perforate the
bark, and run to the scales and leaves. All the vessels
of the wood are dotted. Some of the roots were cut off,
but they entirely agreed in structure with the stem. In
my ' Icon. Sel. Anat. Bot./ vol. ii, I have already shown
the difference between the structure of Dicotyledons and
the Cycadese ; the vessels do not ascend directly upwards,
as in the Dicotyledons, and they traverse all parts of the
bark towards the leaves and scales, which is only the case
in Dicotyledons at individual buds. More recently I
have endeavoured to show, in a short memoir read before
the Academy of Sciences, that the scales are in fact
leaves, and that the so-called leaves are branches. This
renders the germination, especially, very intelligible.

PHYSIOLOGICAL BOTANY.

299

The tap-root, which otherwise is absent in all Monoco-
tyledons, is particularly remarkable.

Observations de Ovulo et Embryonibus Cycadearum.
Auct. T. A. GUIL. MIQUEL. Ann. d. Sc. nat. 3 ser.
vol. iii, p. 193. The following periods of the develop-
ment of the ovule appear to be distinguishable : 1. Before
impregnation, the cellular tissue of the nucleus beneath
the amnios becomes completely absorbed, and leaves a
cavity upon which the amnios lies. On the other hand,
the cavity of the amnios becomes gradually filled with
cellular tissue beginning at the base. 2. This cavity of the
nucleus, which is filled with mucus, then forms a cellular
mass, which does not become connected with the walls of
the cavity, but is inclosed by a membrane ; this is con-
nected with the membrane of the amnios and forms the
albumen. The formation of the albumen does not depend
upon impregnation, for it occurs even in sterile seeds.
The formation of the narrower cavities in the amnios
does not appear to be dependent upon impregnation.
3. Whilst the peculiar cavities are being formed in the
amnios, and the embryoblastanon (of Hartig) is growing
downwards, the entire amnios, excepting its external
membrane, descends into the hollowed apex of the albu-
men which is in progress of formation, and is inclosed
by it, and the apex of the albumen is covered by the outer
open apex of the amnios, as with a cap. 4. The cellular
tissue of the amnios now becomes absorbed, soft ; the
sacs traversing the mucus remain and are covered by a
soft membrane with which they become coherent. 5. As
the embryo enlarges, the embryoblastanon which is re-
flected upwards becomes compressed, the mucous matter
surrounding the sacs dries, and the membrane covering it
disappears, so that when the seeds are ripe, the embryo-
blastanon, with the sacs, are found in the form of an amor-
phous mass, under the persistent membrane of the amnios,
at the point of the root which is forming. The author
then speaks of the anthers of the Cycadeae, and states

300 PHYSIOLOGICAL BOTANY.

that they contain cellula fibroste like other anthers. Lastly,
he characterises the genera Cycas, Macrozamia, Ence-
phalartos, and Zamia, by the form of the embryo. In an
appendix, vol. iv, p. 79, the formation of the albumen
prior to impregnation is confirmed.

On the Plurality and Development of the Embryo in
the Seeds of Conifers. By ROBERT BROWN. Annals of
Natur. History, vol. xiii, p. 369. This paper was read
before the British Association at Edinburgh in 1834 ; it
was published in French in the ' Annal. d. Scienc. Natur/
for October 1843, and then appeared in the above journal.
After alluding to his former views upon the plurality of
the embryos in the Cycadese, which indicates their affinity
with the Coniferse, the author then reports upon his
observations on the seeds of Pinus sylvestris. " The first
and most important change," says he, " is the production
or separation of a distinct body within the nucleus of the
ovule, which before impregnation is a solid uniform sub-
stance. In this stage, the included body or amnios is
slightly concave, and covered with a lacerated cellular
tissue, which either arises from its separation from the
apex of the original nucleus, or of a process which con-
nects it to the apex. Below the concave apex, the amnios
is slightly transparent for about one fourth of its length,
the remaining portion being perfectly opaque ; it consists
of cellular tissue. Before the appearance of the embryos
or the funiculi, the areolse in which they are produced are
visible. These areolse, three or five in number, as ob-
served in May 1827 in the larch, are arranged in a cir-
cular or elliptical series, near the apex, with which they
probably communicate by some points which it is difficult
to distinguish. In Pinus sylvestris^ they were consider-
ably more developed in June or July, from four to six in
number, consisted of conical membranes of a brown colour,
with their apices turned towards the surface, and at the
base seeming to pass gradually into the lighter-coloured
pulpy mass of the amnios. Corresponding and nearly

PHYSIOLOGICAL BOTANY.

301

approximated to each of these conical membranes, a long
funiculus was found ; this was either simple or giving off
a few branches, and generally consisted of four series of
long transparent cells. The upper extremity of each funi-
culus was considerably thicker, of a depressed spherical
form, and each cell contained one of the granules (areolae),
as is frequently observable in the Monocotyledons. In
Pinus pinaster, the funiculus has no appearance of sub-
division, but this finally appears at its ends. By tracing
their development until it is completed, we see that each
of the dark bodies, in which the funiculi terminate, are
embryos in a rudimentary state. Hence the author con-
cludes, that the plurality of the embryos in the order
Coniferae is perfectly constant. In a postscript in 1844,
R. Brown shows, that he had already given the plurality
of the embryos in the Cycadeae in his ' Prodromus Flor.
N. Holland./ but that Dupetit-Thouars had discovered
the principal fact. He then alludes to Schleiden's theory,
and says : " Schleiden ascertained the existence of my
areolae or corpuscles, and denominates them large cells in
the embryo-sac or albumen ; he states, that he has suc-
ceeded in preparing free the whole pollen-tubes from the
nucleary papillae to the bottom of the corpuscles. But if
my observations are correct," adds Brown, " and they
appear to be confirmed by those of Mirbel, the corpuscles
in Pinus are not developed until the following spring or
summer, and hence if Dr. Schleiden's assertion is correct,
the pollen must remain inactive for at least twelve months.
This is not altogether improbable," says Brown, " but
even if it were the case, it would not lead to the adoption
of Schleiden's theory. With respect to the Cycadeae,
adds Brown, under any circumstances it is certain that
the mere enlargement of the fruit, the consolidation of
the albumen, and the complete formation of the corpuscles
in its apex, are wholly independent of male influence ; for
he had seen instances of this in England, when the male
plants of the female Cycadeae which were examined did
not exist in the country.

302 PHYSIOLOGICAL BOTANY.

On the ApocynecB. By ALPHONSE DE CANDOLLE. Ann.
d. Scienc. natur., 3 ser. vol. i, p. 253. This paper is
referred to here, on account of the investigations which it
contains upon the stipules of this plant.

Memoir e sur la Famille des Primulacees. By M. J.
E. DUBY. Geneva, 1844. Germination of the seeds of
Cyclamen, where the large tuber is directly formed, and
the cotyledons are not developed.

Researches upon the Development and the Structure
of the PlantaginecB and Plumbagineae. By M. F. M.
BARNE'OUD. Compt. rend. 1844, ii, p. 262. I. Planta-
ginece. When the flowers are examined in their earliest
state, the development is found to occur from without
inwards, in opposition to Schleiden's theory. The flowers
at first consist of four tubercles, which have exactly the
form and structure of the anthers ; each also is furnished
with a bundle of spiral vessels, and they unite into a tube.
Hence the flower is a tube which supports the stamens,
as in the Gomphrenece and Acliyrantliea. The margins
of the valves of the ovary are at first some distance apart
from each other, and continue to approximate, but never
come completely into contact, hence there are no axile
bodies in the ovary of this order.

II. Plumbaginece. The symmetry in this tribe appears
to be anomalous, because there is one row of stamens
which are opposite the petals. But the author has
discovered the rudiments of stamens in Plumbago mi-
crantha ; they, however, soon disappear, so that the row
of large stamens then forms the rule.

Observations upon the Genus Aponogeton and upon its
Natural Affinities. By J. E. PLANCHON. Ann. d. Scienc.
natur., 3 ser., vol. i, 107. Also Compt. rend., 1844,
ii, 227. The author correctly separates this genus from
the Saurureae, and approximates it to the Alismacese. The
germination is very accurately described here. A single

PHYSIOLOGICAL BOTANY. 303

cotyledon with two rudimentary radicles, and a plumule
issuing from a fissure, the leaves not being sheathing.
But the figures themselves must be referred to.

On the Anatomy of Aldrovanda vesiculosa. By Pro-
fessor PARLATORE. Giorn. Enciclop, i, 237. Compt.
rend. 1844, i, 998. A minute description of this plant,
which is known by its vesicles (ampulla*), which are in
fact leaf -blades, as in Utricularia. The following is
worthy of remark : " The part situated upon the ampullae
is composed of somewhat elongated irregular cells, and
exhibits peculiar bodies, such I have never seen before,
and which, I think, have never been noticed by any
botanist. These corpuscles, which are very numerous
and close together, resemble a small open pair of scissors,
as four arms are easily recognised in them, which are
connected in the centre by a kind of knot." I have found
these scissor-like plates.

On the Surface of the Stem and the Contents of the
Cells of the Pith of Nuphar lutea Sm. By J. MUNTER.
Bot. Zeit., 1845, 505. The author has made the remark-
able observation that the pits in the stem (cormus),
beneath the scars of the petioles, are caused by roots
which separate spontaneously ; a phenomenon which has
not previously been observed in the vegetable kingdom.
In the cells of the pith, he detected the same forms of
starch which he had previously observed in Alstrcemeria.

Researches upon the Structure and Development of
Nuphar lutea. By M. AUG. TRECUL. Ann. d. Scienc.
nat., 3 ser. vol. iv, 286. An anatomical memoir, much
of the contents of which is very excellent. The author's
views are rather too special in investigations of this kind,
and it would occupy far too much space to criticise this
memoir.

On Clandestina Europcea. By M. DUCHARTRE. Compt.

304 PHYSIOLOGICAL BOTANY.

rend. 1844, i, p. 93. This paper contains a statement to
the effect, that stomata exist upon the leaves and young
stems of Clandestina Europ&a. There is a Report upon
the complete Memoir in the ' Compt. rend/ 1845, i,
p. 1268. The so-called etui medullaire is absent in Clan-
destina, neither do medullary rays exist.

Note upon Orobanche Erynyii, Vaucli. By M. P. Du-
CHARTRE. Ann. des Scienc. nat., 3 ser. vol. iv, p. 74.
This plant is furnished with stomata. On the absence
of medullary rays.

In the second part of my ' Lectures on Botany/ many
details regarding the structure of the stem are contained,
which, as far as I am aware, are not stated elsewhere.
It was my wish not to have spoken of them when treat-
ing of the stem in this Report, because I have been found
fault with for having alluded too frequently to myself.
But I was very anxious that many points, as e. g. the
wedges in the wood and in the bark, and the difference
between them, and their difference from the medullary
rays, should not be overlooked.

FERNS. MOSSES. LICHENS. ALG.E. FUNGI.

New Species of the Genus Isoetes, from Algeria. De-
scribed by BORY ST. VINCENT. Compt. rend. 1844,
i, 1167. They consist in addition to Isoetes Delilii, of
Is. longissima, from bogs, and /. Duriei and Hystrix,
terrestrial species. Bory quotes Cl. Richard as having
stated that the Isoetese should constitute a separate
natural order; this would also apply to Salvinia and
Pilularia. Roeper in his Flora of Mecklenburgh, pt. i,
1843, which contains much excellent matter, particularly
special remarks upon Ferns, has censured me for arranging
the Lypodiacese among the Ferns, and the same may be
said of the Equisetacese. But I should wish the whole
to be united into one class, because the orders composing

PHYSIOLOGICAL BOTANY. 305

it are, as it were, the representatives of a Flora of the
former world at a certain period. Moreover they bear
the characters of these plants ; everywhere we find im-
perfectly developed structures, not yet separated ; the
internal structure of the stem of the Lycopodiaceae is that
of a root, the frond is both a stem and a leaf; the male
and female sexual organs are still conjoined in the
Salviniacese, &c.

MoveaUe Spiral Threads in Ferns. By C. NAGELI.
Zeitschr. f. wissenschaft. Botanik, Hft. i, p. 169. On
the under surface of the germinal leaf, at its margin, more
rarely also upon the upper surface, occur glandular organs.
They frequently appear to consist of a single cell only ;
we generally recognise that it is a sac, composed of a
single layer of cells. This sac is filled with apparently
granular and opaque contents ; it bursts at the top, and
allows a number of small round cellules to escape ; these
cellules move actively in water. Each contains a spiral
thread, which is set free by the rupture of the membrane
of the cellule, and then exhibits the same movements as
the seminal filaments of the Mosses, Liverworts, and
Charas. An interesting addition to the observations
which have already been made upon these Entophytes.

Lamella of the Leaves of Mosses. By C. MULLER.
Linnsea, vol. xviii, p. 99. The elevated ridges which
are found upon the upper surface of the leaves of several
Mosses on the side of, and in connexion with the nerves,
were first accurately described by Treviranus, and are
again minutely treated of in this paper. They consist of
a row of remarkable cells ; they do not seem to be of any
use ; they are connected with formative principle,

On the Development of the Sporidia in the Capsules of
Mosses. Dissert. Inaug. scr. Bo. Jung. Scato. GEORG.
LANTZIUS BENING. Gott. 1844. An excellent contribu-
tion to the investigation of Mosses, The arrangement of

306 PHYSIOLOGICAL BOTANY.

the sporidia in groups of four, in the form of a tetractys,
has often been observed since Mohl described it, but their
formation has never been traced so accurately from its
earliest origin, as by the author. His observations show
that the development of sporidia in the capsules of the
Mosses, is effected by a frequently repeated formation of
cells within cells, which takes place four times in Poly-
trichum, three times in Hypnum, but only twice in several
other Mosses. It could not be satisfactorily determined
whether the separation of the cells was produced by
internal walls, or the granular matter; probably both
contribute to it ; a wall is formed on the separation of the
nucleus. The formation of cells within cells never occurs
during the simple course of vegetation, but this is the case
where vegetation returns to generation.

History of the Growth of Mosses and Liverworts. By
C. NAGELI. Zeitschrift f. wissenschaft. Bot., pt. 2, 138.
The history of the growth of Echinometrium and seve-
ral other Liverworts does not admit of an abstract.

G. METTENIUS, Contribution to the History of the
.Development of the Mov cable Spiral Fibres of Char a
hispida. Bot. Zeit. 1845, p. 17. " The two antennae
which Thuret constantly observed, I have not been able
to find/ 5 says the author ; I have seen them, and this
entirely alters the matter.

Identity of the filamentous and mucoid Conferva. By
Dr. SCHAFFNER. Flora, 1844, p. 568. Rendered pro-
bable, but not proved. These minute objects certainly
much resemble each other ; they are Fungi, not Confervse.

Caulerpa prolifera Ag. By C. NAGELI. Zeitschrift f.
wissenschaft. Botanik, pt. i, p. 131. History of the
Development of Delesseria Hypoglossum. By the same
author. (Ibid. pt. ii, p. 121.)

PHYSIOLOGICAL BOTANY. 307

As regards the Algae, descriptive coincides so closely
with physiological botany, that the two must be arranged
near each other. We find, in the ' Annals of Natural
History/ vol. xiii, p. 375, RALFS, On British Desmidete ;
vol. xiv, p. 240, DAY. LANDSBOROUGH, On the Fructifi-
cation of Gloiosiphonia Capillaris ; ibid. pp. 256, 361,
On British Desmidea continued.

The following papers are important as regards the
description of the Algae : On the Structure of Ctenodus,
Delisea, and Lenormandia. By M. MONTAGNE. Annal. d.
Sc. nat., 38, vol. i, p. 151. Note upon some Alga with
reticulated Fronds. By M. J. DECAISNE. Ibid. vol. ii,
p. 233. Note tipon the mode of Reproduction of Nostoc
verrucosum. By M. GUST. THURET. Ibid. p. 319. Ob-
servations on the Tetraspora. By M. M. CROUAN. Ibid,
p. 365.

On the Colour of the Red Sea. By M. MONTAGNE.
Ann. des Sc. nat., vol. ii, p. 332. Account of it, and
confirmation of Ehrenberg's earlier investigations. It is
produced by an Alga, which Ehrenberg denominated
Trichodesmium erythraceum.

On the genus Ceramium and some of its Species. By
Prof. MENEGHINI. Giorn. Encicl., i, p. 178. A cri-
ticism of Kiitzing's Memoir in the 'Linnaea/ vol. xv.
Professor Meneghini works principally at the Algae.

On the History of the Development of Chara. By
C. MULLER. Botan. Zeit. 1845, p. 393 et seq. The
formation of cells has been treated of at some length
above. This paper principally consists of a history of
the development of the cells of Char a from cytoblasts.

On Hamatococcus pluvialis. By J. V. FLOTOW. In
the Memoirs of the Royal Academy of Naturalists, b. xii,
sect. 2, p. 413. An important memoir. The author

308 PHYSIOLOGICAL BOTANY.

describes, with great care and accuracy, the transforma-
tions of a small Alga, or a small Infusorium, Hcematococcus
pluvialis, into the most varied forms. A red matter was
first found in the rain-water from a surface of granite
(at Hirschberg) ; this consisted of extremely delicate
globular, shining vesicles, filled with a mass which, in its
moist state, was granular, and of a carmine red colour ;
when dried upon paper, it was of a vermilion red colour.
These granules not only experienced a change of colour,
becoming greenish after some time, but at the end of
September and the commencement of October, move-
ments commenced in the granules. These consisted of
1, Motion in the direction of a curve (longitudinal mo-
tion) ; 2, Rising and sinking in serpentine lines ; 3, A
rotatory motion. Water removed from the above cavity
was then examined from time to time, and the altered
forms studied, examined, and described with extraordinary
accuracy. On the 30th of November filaments began to
be formed ; on the 13th of December he examined some
of the rain-water which had been taken on the 9th of
October, and subsequently kept in a warm room; he
then found an infusorium, Astasia pluvialis, in it, which
is nearly related to A. nivalis Shuttlew. I cannot dispel
the idea, says he, that this Astasia arose from the Haema-
tococcus, and that it is only a higher stage of development.
Its agreement in form and contents with the Hteinato-
coccus globules themselves, the number of intermediate
forms, composed of perfectly round figures, which from
the first become slightly oval or ovale, afterwards more
and more so, and sometimes are free from tubercles,
at others bear them, almost entirely preclude the for-
mation of an absolute line of demarcation between the
phytonomic or infusorial animated individuals. This
Astasia pluvialis is not seen to be produced in any kinds
of infusion which do not contain Htematococcus pluvialis,
the presence of which is a necessary condition to its for-
mation. Moreover, a constant reciprocity is observed to
exist between the two ; the Astasite increase by sub-

PHYSIOLOGICAL BOTANY. 309

division, but their embryo in part again assumes the form
of Haematococcus. I must at least assume this to be the
case. Thus the author saw the Htematococcus increase
in vessels which were set aside, and attach itself to their
sides. He also perceived infusiorially-animated indi-
viduals among them ; but he never saw the Hcematococ-
cus, which was then at rest, subdivide. He separated
the Hcematococcus, and then found that every change of
temperature, or of the water, provided the latter was
pure, and the globules had acquired maturity, produced
an alteration in the H.pluvialis. Experiments detailed
with the same extreme minuteness follow the observations.
They were performed with infusions of Hamatococcus,
and contain a number of very remarkable observations
upon the various forms which this being produces ; but
we must refer the reader to the memoir itself. The
author is uncertain whether he should refer Htematococcus
to the animal or the vegetable kingdom, but concludes in
favour of the latter opinion.

The same subject is treated in a shorter paper : On the
Conversion of Infusoria into the loiver Forms of the Algce.
By Dr. FR. TRAUG. KUTZING. Nordhausen, 1844, 4.
After some historical remarks, in which Flotow's memoir
is also quoted, the author communicates some observa-
tions, the result of which is, that Chlamidomonas pulvis-
culus is susceptible of numerous changes ; that a decided
species of Algae, Stygeoclonium stellare, is developed from
it, but that other formations also emanate from it, which
likewise decidedly have the characters of the Algae,
although from their external form they might in part lay
claim to being considered as infusorial forms in a state
of rest.

In conclusion, says the author : " The natural history
of organisms has hitherto been treated of by two methods,
according as the object is considered as perfected, or in a
transient state. Linnaeus may be regarded as the dis-
coverer of the defining method in natural history, and

310 PHYSIOLOGICAL BOTANY.

Gothe as the discoverer of the exponential method." Why
does the author, who is remarkably clear-headed, make
use of such expressions as these ? Can a body be con-
sidered as in a state of formation when we are ignorant
of what it is destined to become ? Must we not always
commence with the defining method, and then pass to
the exponential ? Have not all natural philosophers done
so ? Were not the species of frogs and salamanders first
determined ? Was this not necessary to avoid confusing
the metamorphoses which they undergo in their early
stages ? Again says the author : " Some of our philoso-
phers assert that it is necessary for us to assume well-
defined limits between plants and animals, since without
this assumption science would degenerate into fantastic
mysticism." Now this is not exactly the case.

But when I distinguish plants from animals, I must
know by what means I do so. Ehrenberg adopts the
character of an animal originally proposed by Blumenbach,
the presence of a cavity (stomach), from which the
entire being is nourished. This is not the place for dis-
cussing whether this is correct or not. Ehrenberg would
ask, has Hcematococcm a stomach ? No : then it is not
an infusorial animal, but a portion, the seed of one of
the Algae, which must undergo several metamorphoses
before it is perfectly developed. For the same reason,
the spore of Ectosperma (Faucheria), with its cilia, is not
an animal, but, as Unger correctly states, in a state of
transition to the animal condition ; that is, as far as we
can judge from its possessing motion. Observations
and experiments, such as those detailed by Kiitzing and
Elotow, are of great importance ; but I should prefer a
somewhat simpler path than the latter has followed.

On the Cells containing Spiral Fibres in Fungi. Bot.
Zeit., 1844, p. 369. The author, Professor von Schlech-
tendal, after reporting what has already been stated
upon this point by Roman. Hedwig, and subsequently
Corda, details his own observations upon some dried

PHYSIOLOGICAL BOTANY. 311

Trichise. The cells are either not very long, and in that
case pointed at both ends and contain few spirals, by
means of which the walls of the cells are, as it were,
expanded ; or the cells are very long, divaricately rami-
fied, and intricately interlaced with each other. More-
over, the spores have always a larger diameter than the
cells containing the spiral fibres, which require high mag-
nifying powers for their observation.

On Lanosa Nivalis Frs. By Professor UNGER, of
Gratz. Bot. Zeit., 1844, p. 369. This remarkable white
filamentous Fungus, the reddish separating sporidia of
which were first described by the author, having only
been mentioned by Fries and Corda, occurred in large
quantity beneath the melting snow at Gratz, at the end
of February and the beginning of March. The author
ascribes the sudden vegetation of this Fungus to the cir-
cumstance, that notwithstanding the heavy fall of snow
in January and February, the ground was still not frozen.

A sick woman, who was principally suffering from
difficulty of deglutition, vomited some sporidia of a fungus;
these were sometimes arranged in a moniliform manner.
H. Gruby satisfied himself, that all the nourishment
she took was fresh and good. Compt. rend., 1844, i,
p. 586.

Occurrence of Cysts, containing Filamentous Fungi, in
the Internal Ear of a Young Woman. By Professor
MEYER. Muller's Archiv, 1844, p. 404. The descrip-
tion and figure distinctly show that this was Mucor
Mucedo.

The Report of the Swedish Academy of Sciences for
1844, contains an instance of AcJilya prolifera prpducing
the death of fish. My friend Lichten stein sent me a
small Cyprinus Alburnus, the mouth of which was en-
tirely closed by the growth of Acklya prolifera. Almost

312 PHYSIOLOGICAL BOTANY.

all the fish in the pond had been destroyed by this growth.
On examining it, I found that the difference stated to
exist between Aclilya and Saprolegnia is incorrect, for
several filaments had transverse walls, many not.

Identity of the Mucoid and Filamentous Conferva.
By Dr. SCHAFFNER. Flora, 1844, p. 567 ; also Flora,
1845, p. 501. In an appendix to his remarks upon the
Mucoid Confervse, Dr. Schaffner states that this plant is
Byssocladium fenestrale. He also found this Byssocla-
dimn in the expectoration of a patient suffering from
pulmonary tubercle. The pulverulent crusts of Porrigo
leprosa, as well as those of scrofulous scald-head, also
consist of a variety of Byssocladium fenestrale. ' Flora/
1845, p. 501. There is no doubt that many of these
Fungi are still imperfectly developed. How many Eld-
zomorphete do not consist of the thallus of Merulius
(Xylophagm vastator), the dry-rot ! Still more remark-
able are the filaments which are developed in syrup pre-
serves, and even in solutions of the tartrates. They must
be allowed to grow for a long time before we can recognise
them as belonging to Penicillium glaucum. I cannot
too strongly recommend that these Fungi should be left
undisturbed, so as to allow of their true fructification
being discovered. Hitherto, there has been much con-
fusion on this subject. Nothing decidedly botanical has
been published upon the Fungi found in fermenting
fluids.

MONSTROSITIES.

On some Abnormally-developed Leaves. By the Editor,
VON SCHLECHTENDAL. Bot. Zeit., 1844, p. 441, 457.
Contains a notice of some abnormally-developed leaves
which were observed; hence it does not permit of an
extract.

The monstrosities described by Kirschleger would re-
quire to_be detailed in full. Vide ' Flora/ 1844, pp. 129,

PHYSIOLOGICAL BOTANY. 313

566, 728 ; 1845, pp. 402, 613. I may mention, also, the
" Antholyses" of Valentin, N. Act. Acad. Leop. 18, i,
223 ; Duchartre's two cases in Ann. d. Scienc. Natur.
3 ser., vol. i, p. 292 ; and those of Cappari, Giorn.
Encicl., vol. ii, p. 261.

Upon a very Rare and Special Ramification of Yucca
Aloifolia L. By ANTONIO PRESTANDREA, of Messina.
Messina, 1845, p. 8. Such ramifications are by no
means rare, especially in warm climates. I have fre-
quently observed the ramification of the scape of Agave
Americana below the inflorescence, in Messina.

A Monstrosity of Primula Sinensis, in which a cup-
shaped body upon which the naked placenta was situated,
occurred upon the style, is very remarkable. Babington,
Ann. of Nat. Hist., xiii, p. 464. Dr. Dickie describes
several monstrosities in Gentiana campestris, ibid, xv, 87.

Monstrosities in Digitalis purpurea through several
generations, observed by Vrolik. Flora, 1844, i.

Professor von Schlechtendal's Remarks upon a Mon-
strosity of the Capsule of Papaver Somniferum, are also
extremely interesting. Bot. Zeit., 1845, p. 6.

REPORT'

ON THE PROGRESS OF

GEOGRAPHICAL BOTANY,

DURING THE YEAR, 1844.

BY DR. A. GRISEBACH,

EXTRAORDINARY PROFESSOR IN THE UNIVERSITY OF GOTTINGElf.

GEOGRAPHICAL BOTANY.

LN the first volume of the ' Physical Atlas' of Berghaus,
which is now completed, six sheets are devoted to graphic
representations in the department of Botanical Geography.
The first sheet, entitled ' Outlines/ forms a sequel to the
works of Humboldt and Schouw, and refers principally
to the geographic subdivision of vegetable formations ; in
the vertical direction it illustrates the serial gradations
of the regions, whilst in the horizontal direction it shows
the areal boundaries of the natural Floras. This repre-
sentation, however, appeared as early as 1838, and on
future revision would require considerable improvements.
The second sheet, which treats of the Districts in which
the most important products of culture are distributed, is
of greater interest. Its design consists in an attempt to
subdivide the province of agriculture throughout the en-
tire inhabited surface of the earth, according to the kinds
of Cerealia which predominate, whence general relations
are found between the climate and the productive power
of different countries. In the Old World the author dis-
tinguishes the following zones between the polar limits to
agriculture and the equator.

1. Zone of barley and rye. It might with propriety
be called the zone of the summer Cerealia, inasmuch as
the duration of the winter is the most important condi-
tion which prevents the culture of the more productive

318 GEOGRAPHICAL BOTANY.

and certain winter corn. In this more comprehensive
point of view, the separate denomination of the South of
Scandinavia as the district of the exclusive cultivation of
rye, and of Scotland as that of barley, disappears, as
circumstances not founded upon climatic conditions.

2. Zone of rye and wheat. This is considered as
extending southwards to about the fiftieth degree of
latitude, or as far as the polar limits of the cultivation of
the vine.

3. Zone of wheat. To this, those parts of Europe
and Western Asia belong which lie south of the fiftieth
degree. In several districts maize is cultivated as well as
wheat.

4. Zone of rice and wheat in those provinces which
are subject to the influence of tropical seasons. In tro-
pical Western Africa rice and maize occupy the place of
the former.

In America, where these relations are modified by the
greater extent to which maize is cultivated, Berghaus
distinguishes the following zones : rye, wheat, and barley
(i. e. summer Cerealia) ; rye and maize ; wheat and
maize ; wheat ; in the tropical zone maize is the principal
cereal grain. With these sketches the author has com-
bined indications of the distribution of other nutritive
plants, and has illustrated, in separate charts, the districts
in which the most important plants of commerce are
produced. The two following sheets contain the statistical
numerical proportions of the Flora of Europe, which, not
being susceptible of tabular arrangement, and being sub-
ject to very important differences in the views taken of
the definition of the species and botanical groups, were
not adapted, in the present state of botanical geography,
to graphic representation. Although the same applies
still more to the last sheet upon Germany, which appeared
in 1841, nevertheless the review of the polar and
equatorial limits of numerous woody and cultivated
plants in Europe, deserves great praise, inasmuch as the
observations made use of in it have appeared to us,

GEOGRAPHICAL BOTANY. 319

on the frequent use of this chart, very numerous.
Moreover, many of the sheets intended to illustrate
meteorological relations appear indispensable also to the
botanist.

M. Romerhas commenced the publication of a Memoir
entitled ' Botanical Geography and Geographical Botany/
which treats of the subdivision of the surface of the earth
into natural Floras. (Liidde Zeitschr. fur vergl. Erdkunde,
Bd. iii, pp. 527-534.)

A paper by E. Fries, entitled " The Native Land
of Plants," in his peculiar style, the special interest of
which is confined to the Swedish public, but also fre-
quently touches acutely more general questions, treats of
different botanico-geographical subjects, especially of the
native country of the so-called ruderal plants. (Botaniska
Utflygta, Bd. i, pp. 229-328, translated in Hornschuch's
Archiv Skandinav. Beitrage zur Naturgesch. Bd. i,
H. 3.) The original native country of many cultivated
plants cannot now be determined by empirical proof, but
only by rational investigation. Thus rape is no longer
met with in its wild state, but when we adduce proof
from all extra-European countries that it is not indigenous
to them, we must conclude that it is of European
origin, although its wild state has disappeared through
cultivation. Many plants have been extirpated by use ;
this is now gradually taking place with Gentiana lutea,
in the Alps, and Inula Helenium in the west of Sweden.
The contact of Nature with man exerts no less a modify-
ing influence upon the vegetable kingdom than upon the
animal creation. The original vegetation of a country
must in general, therefore, be regarded as more rich in
species, and in this manner in Sweden and Germany,
even under our own eyes, the localities of rare plants are
disappearing one after the other, as e. g. of Trapa, Xan-
thium, and Stipa.

320 GEOGRAPHICAL BOTANY.

The excellent work of A. Wagner, on the ' Geographi-
cal Distribution of the Mammalia/ (Abhandlungen der
mathem. physik. Klasse der Bairischen Akad. Bd. iv),
which belongs to an allied province, but was not designed
without regard to the geographical relations of other
organisms, must not be passed over here without notice.
The question of the original native country of the various
organisms is acutely investigated by the author, and it is
found that the distribution of animals, as of plants, can-
not be satisfactorily explained by the climatic and local
conditions of their existence, but that the most rigid facts,
together with the physical relations at present in existence,
point to other, perhaps historical causes, with which we
are at present unacquainted, and which the author con-
siders as the effects of a general order of the creation,
which ought, however, rather to be kept in view by us as
objects worthy of future investigation. From the observa-
tions, in Belgium, upon the Periodical Phenomena of
Vegetation, published by Quetelet, and mentioned in the
previous Yearly tteport, the following brief extract, con-
taining the period of the appearance and fall of the leaves,
in the year 1841, of some generally diffused woody plants,
may be of use in the determination of the Phyto-isotherms
of Northern Europe ; and for this purpose it will be con-
joined with some observations simultaneously made by
Hartmann, in Gefle (60 N. 1.) (Bot. Notis. 1842.)

GEOGRAPHICAL BOTANY.

321

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-I "51

322 GEOGRAPHICAL BOTANY.

Of monographs upon individual groups of plants, in
which attention is paid to their geographical distribution,
published during the past year, the following require
mention: Parlatore on the Fumariacese (Giornale Botan.
Ital., i, p. 97 et seq.); v. Martins, on the Erythroxylaceae
(Bairische Abhandl., iii, pp. 325-32); Lomler, on the
Distribution of the Coniferse (Ratisbon Mora, 1844,
pp. 440-3).

Fumariacece. Only 1 3 species ; these are distributed
throughout both temperate zones, for the most part,
indeed, secondarily transferred from one region to the
other. With the exception of the Cape Discocapnus, they
all grow in the South of Europe, between the 34th and
40th degrees of latitude, and diminish so rapidly from
this zone in both meridional directions, that beyond the
50th degree, 3 species only are met with; a statement
which, however, is not correct as regards Germany.
Spain contains several endemic forms.

Erytliroxylacea. Of 58 species of the genus Ery-
tliroxylon, Brazil contains 29; the West In dies, 8; Guiana,?;
Columbia, 4 ; and Mexico and Peru, one each ; hence
tropical America contains 50 altogether : 5 species grow
in Madagascar and the Mauritius, single representatives
at the Cape, in the East Indies, and on the north coast
of New Holland. In America the district of their distri-
bution extends from the tropic of Cancer to that of Capri-
corn, in the Old World, from 15N. lat. to 30 S. lat.

Conifer a. Lomler enumerates only 208 species. Of
these, he calculated that 165 exist in the northern and
51 in the southern hemisphere ; moreover, there are 22
in Europe, 87 in Asia, 16 in Africa, 83 in America, and
35 in Australia; lastly, 24 in the tropic zone, 159 in the
north temperate, and 33 in the south temperate zone.
These statements can only be regarded as preliminary
steps to our knowledge on this point.

GEOGRAPHICAL BOTANY. 323

I. EUROPE.

A work, containing copper-plate engravings of the
plants of Russia, has been begun by Trautvetter (Planta-
rum Imagines et Descriptions . Monachii, 1844, 4 fasc.,
1-4 ; at present 20 plates). Also a continuation of the
old ' Bieberstein Centuries' (M. de Bieberstein Centuria
Plantarum Rossise Meridionalis Iconibus illustrata ; Pt.
ii, Dec. 1-3. Petropoli, 1844,) has been commenced
in St. Petersburg. Engelmann has published a paper
upon the Genera of Plants found in the Russian provinces
of the Baltic (Genera Plantarum, or the Genera of Plants
growing wild in Esthonia, Livonia, and Courland ;
Mitau, 1844-8).

A. E. C. v. Fischer has written upon the botanical re-
lations of Southern and Central Lithuania, especially in
the circle of Sluzk (Mittheilungen d. Natur. GeseUschaft
zu Bern, for the years 1843-4; Bern, 8). In the im-
mediate neighbourhood of Sluzk, in the district of the
source of the Niemen and several tributary streams of the
Dnieper, the author only found about 600 Phanerogamia,
a catalogue of which he gives, with remarks upon their
statistics. In these districts, heathy plains, overgrown
with Calluna (together with Juniperus and Genista tine-
toria) are still common. Dwarf underwood, consisting of
the oak (Quercus pedunculata) covers large spaces, and
stamps the physiognomy of Lithuania towards the western
districts of the Baltic plain. In moist low grounds Salix
angustifolia and livida predominate. The large forests
consist of pines or fir trees ; the truly foliaceous trees,
which are less common, are mostly birch, and in Polesia,
the oak, which grows mixed with the birch, poplar, moun-
tain-ash, &c. The following may be mentioned as geo-
graphically characteristic species: Thalictrum aquilegifolium
L., simplex L., and anyustifolium Jacq., Anemone patens~L.,

324 GEOGRAPHICAL BOTANY.

Viola stricta Horn., DiantJtus arenarius L., Euonymus
verrucosus Scop., Trifolium lupinaster L. (in pinetis sic-
cioribus raro), Spiraea Aruncus L., Geum strictum Ait.,
Polentilla norvegica L., Agrimonia pilosa Led., Saxifraga
Hirculus L., Cnidium venosum Kch., ChceropJiyllum
aromaticum L., Inula Helenium L. (in sylvis udis),
/. hirta L., Cirsium rivulare Kch., Andromeda calycu-
lata L., Pyrola media Sw., Polemonium c&ruleum L.,
Pulmonaria azurea Bess., Pedicularis Sceptrum L.,
DracocepJtalum Huyscliiana L., Melittis Melissopliyllum L.,
Amaranthus sylvestris Desf., Thesium ebracteatum Hayn.,
Euphorbia virgata Kit., Salix nigricans Fr., livida Wahlb.
(depressa Fr.), myrtilloides L., versifolia Wahlb., lap-
ponum L., BetulafruticosaYa)k., Typha pendula nov. sp.,*
Malaxis monopJiyllos Sw., Cypripedium Calceolus L.,
Gladiolus imbricatus L., Fritillaria sp., Veratrum
Lobelianum Bernh., Tofieldia calyculata Wahlb., Car ex
divulsa Good., pilosa Scop., Hierochloa odorata Wahlb.,
Calamagrostis stricta Spr.

Wahlberg has published some remarks upon the plants
of Quickjock in Swedish Lapland (Ofversigt af Kongl.
Vetenskabs-Akademieens ForhandL, 1844, p. 23). Rubus
castoreus Laestad. is a bastard of E. articus and saxatilis
occurring in two forms.

Lindblom has published some observations upon the
Botanical Relations of Norway (Bot. Notiser., 1842-3.)
In the outset, we meet with the unfounded assertion, that
in most of the regions of the coast of Norway, Alpine
plants extend as low down as the level of the sea ; an
occurrence which is limited to individual species only,
and may be compared to the growth of Alpine plants on
the Isarkies, near Munich. This statement, made by
Lindblom, is one of those erroneous generalizations, bor-
rowed by one person from another. Alpine plants do
not occur in Norway below the limit of trees, any more

* T. spicis cylindricis, masc. et fcem. contiguis, foliis plains linearibus
culmo longioribus pendulis. (An. T. Shuttleworthii, Kch. ?)

GEOGRAPHICAL BOTANY. 325

than in the Alps. Then follow observations upon the
limits at which plants occur in the direction from west to
east, to which scientific value must be attributed, on ac-
count of the climatic contrasts between the internal dis-
tricts and the western coast of the south of Norway.

a. Plants of the western coast, which, according to
Lindblom, are not found in the inner district. (The
polar limit of their distribution is expressed numerically
according to the degree of latitude, their occurrence in
Sweden is inclosed in parentheses).

Fumaria capreolata. 59. Erica cinerea. 62^.

Hypericum pulchrum. 63^. Pyrola media. 61.

montamtm. ( Bohuslan.)

Vaerdal in Trondjem. Lysimachia nemorum.^.

Vida orobus. 62|, i. e. the limit (Schonen.)

of the occurrence of the oak. Primula acaulis. 63(=).

Sanguisorba officinalis. 00. Digitalis purpurea. 63.

(Isld. Gottland.) ( Bohuslan.)

Bunium Jlexuosum. 63. Lamium intermedium. 61.

Myrrkis odorata. 63. Teucrium Scorodoma. 59.
Chryospleniumoppositifolium. 62|. Luzula maxima. 68.

Rosa pimpinellifolia. 60. Carex binervis. 63.

Ilex aquifolium. 62|. salina. 70.

(Bohuslan.) maritima. 70.

Gali urn saxatile. 62^. Air a prtecox. 62^.

(South of Sweden.) ( Bohuslan.)

Centaurea nigra. Bromus tectorum. 61.

Snaasen in Trondjem. Brachypodium gracile. 62^.
Hypochteris radicata. 62^.

( Bohuslan.)

b. Plants belonging to the western coast, which occur
only on the southern coast, e. g. at Christiania, or in the
valleys of the Fjeldplateaux, but not in the true inner
districts of the south of Norway.

Arabia petrow.W . Hedera helix. 60^.

Rosa pomifeta. 63. Lonicera periclymenum.

(South of Sweden.) Valderhong in Trondjem.

Sorbus aria.Q3%. ( - Bohuslan.)

( Bohuslan.) Sambucus rdgra.
Sorbus hybrida.W? (?) Valderhong. ( Bohuslan.)

(Gottland.) Gentiana purpurea.W\.

326 GEOGRAPHICAL BOTANY.

Mentha sativa.. Quercus Robur. 62.

(South of Sweden.) According to Blom, 63.

Fagus sylvatica. 61. (South of Sweden.)

( Bohuslan.) Allium ursinum. 63.

( Bohuslan.)

c. Inland plants of the eastern districts of southern
Norway, which are absent on the western coast. (Ex-
cluding those of the Ejeld.)

Pulsatilla vernalis. Dracocephalum Ruyschiana.

Trottius EuroptBus. Thymus Chamaedrys.

Berberis vulgaris. Pedicularis Sceptrum.

Ledum rupestre. amygdalina.

Galium trifidum. Carex capital a.
Hieracium cymosum. parallela.

Pyrola chlorantha.

d. Plants of the Eastern Fjeld, principally observed on
the Dovre-fjeld, but not found on the western coast (some
species which I myself found at Hardanger, and which
are therefore more widely distributed, are omitted in this
list, viz. Aconitum septentrionale, Draba hirta, Gentiana
nivalis, and Salix arbuscula).

Ranunculus hyperborem. Saxifraga controversa.

Lychnis apetala. Primula stricta.

Alsine hirta. Gentiana tenella.

Oxytropis lapponica. Koenigia Islandica.

Phaca oroboides. Junctus arcticus.

frigida. Kobresia caricina.

Potentilla nivea. Elyna spicata.

Saxifraga cernua. Carex microglochin.

A remarkable peculiarity of the highlands of Norway,
and which is not merely indicated, but satisfactorily
established by this catalogue, yet cannot be explained by
means of the variations of climate pointed out above,
consists in the fact that the Alpine vegetation appears to
attain its maximum, as regards the number of species, on
the Dovre mountains, and that it diminishes from this
locality both towards the west and the south. Moreover
in these directions the individual numbers of many cha-

GEOGRAPHICAL BOTANY. 327

racteristic species also become less, the Fjeldplateau
gradually assuming the condition of a steppe. In this
respect, Lin dblom's observations on the desert of the By gle-
an d Hekle-Fjelds, or the most southern part of the high-
lands, which were made many years ago, but are again
brought forward in the present memoir, are instructive.
The predominating plants of some tracts in this part,
e. g. between Siredal and Lysefjord, are Molinia ccerulea
and Solidago virgaurea, and these displace all others.
The alpine plants of this region, as shown by the following
list of them, also grow in Hardanger, and do not resemble
those of the Brocken or the Sudeten, to which, among the
whole of the Scandinavian mountains they are most nearly
situated.

Arabia alpina ; Cardamine bellidifolia.

Silene acaulis; Lychnis alpina; Stellaria alpestns; Cerastium trigynum

alpinum ; Sagina Linnaei.
Epilobium alpinum, alsinifolinm.
Dry as octopetala ; Potent ilia maculata ; Sibbaldia procumbens ; Alchemilla

alpina.

Rhodiola rosea.

Sax if rag a Cotyledon, stellaris, aizoides, rivularis, oppositifolia, nivalis.
Sanssurea alpina ; Hieracium aurantiacum, alpinnm.
Phyllodoce taxifolia ; Cassiope hypnoides ; Arctostaphylos alpina ; Loiseleuria

Veronica alpina, saxatilis ; Bartsia alpina.

Oxyria renifurmis.

Salix glauca, Myrsinites, Lapponum, retusa, herbacea.

Detula nana.

Tofieldia borealis.

Jiaicus biglumis, trifidm ; Lnzula arcuata and spicata.

Aira alpina and atropurpurea ; Poa alpina ; Phleum alpinum.

Carex rariflora, pulla, lagopina, rigida, vaginata, atrata, rotundata, capillaris

and alpina ; Eriophorum capitatum.
Lycopodium alpinum.
Polypodium alpestre.

The second section of Lindblom's memoir treats of the
distribution of the Norwegian Ferns, which, according to
theory, ought to be more common on the western coast

328 GEOGRAPHICAL BOTANY.

than in the inland districts, but which, in fact, do not
correspond to this view. The author is certainly of an
opposite opinion, and states that the number of indi-
viduals increases towards the west, which I should much
doubt ; but it is certain that Hymenopliyllum Wilsoni can
alone be considered as an evidence of the marine climate,
whilst the inland country contains five more of the thirty-
three ferns which are here enumerated than the west, viz.
Polypodium calcareum; Aspidium Thelypteris, cristatmn,
montanum, and crenatum Sommf. On the western coast,
Aspidium aculeatum and Asplenium adiantum nigrum,
which are not found in the east, extend as far as
Trondjem, but they must be considered as forms belong-
ing to the south, not to the coast.

I can only refer here to my paper upon Hardanger (see
Wiegm. Arch., p. 1-28) ; still I cannot omit this oppor-
tunity of replying to the editor of the ' Botaniska Notiser'
(see that journal, 1844, appendix, p. 64), that the beech
is certainly cultivated beyond Christiansund. Blom, whose
authority is Blytt, makes this statement. (Das Konigreich
Norwegen. Leipz., 1843, p. 48.) I did not say that it
grew wild there, as Lindblom has erroneously stated in
his translation, and the only object I then had in view,
was to show how far north the climate was suitable to
the growth of that tree. I found single specimens of
Helianthemum alpestre on rocks near the herdsmen's hut
Oppedals-Stolen, and have given specimens of PJtippsia
from the same region to several botanists. However, I
place little value upon these new localities, of which I
had several, and I should consider it as the best recom-
pense for the labour of my memoir, if Lindblom and
other able Scandinavian naturalists, instead of filling
their Journal with unsatisfactory lists of the results of
their excursions, and critical minutiae regarding the dis-
tinction of species and their nomenclature, were also
induced by it to direct their scientific attention more and
more to the conditions of the distribution of plants in the
North of Europe.

Blytt, whose Flora of Norway has long been in prepa-

GEOGRAPHICAL BOTANY. 329

ration, but is still looked for in vain, has published a
Catalogue of the Plants growing wild at Christiania
(Enumeratio Plantarum, quse circa Christianiam sponte
nascuntur, Christiania, 1844, p. 4). It contains 790
vascular plants. Fries has continued the publication of his
Critical Remarks upon Swedish plants and their stations
(Bot. Notis. 1844, p. 1, 49, 75 et seq.) Parts ix and x of
his Normal Herbarium have appeared. Anderson and
Lindblqm have worked at the Alpine Epilobia of Sweden
(id.) Angerstrom has issued some contributions to our
knowledge of the Scandinavian Mosses (Nov. Act. soc.
Upsal. 12, pp. 345-80).

Lindblom's Botaniska Notiser also contains the follow-
ing Memoirs upon the Topography of Swedish plants :
Borgstrom, Contributions to the Flora of Warmeland
(1842) ; Lindgren and Torssell, Mosses of Upsal (1842-3) ;
Forssell, Catalogue of the more rare Plants which occur
in Norrtelge (north-east of Stockholm) (id.); Hofberg,
Localities at Strengnas on the Lake Malern (1842-3) ;
Von Post, Botanical Conditions of the Western Bank of
Lake Malern (1844), of some interest, on account of
the careful observation of the localities in which 480
phanerogamous plants are distributed; Hamnstrom, New
Localities in Nerike (1842) ; Lindgren, Localities at Lake
Wener, with critical remarks (1842-3); Holmgren,
Kalen, and Hamnstrom, Localities in East Gothland
(1841-3); Lagerheiin, the same in West Gothland (1844);
Sieurin, Diary of Travels in North Holland, containing
habitats (id.) ; Lindblom and Borgstrom, Habitats at
Schonen (1843-4). Nyman has published Contributions
to the Flora of Gothland, by which the number of vascular
plants found upon this island is increased to more than
800 (Vetenskaps Akademieens Handlingar for ar 1840,
pp. 123-51). The results of Beurling's voyage now com-
municated in these Memoirs, are confined to lists of
localities, principally in Jemtland ; they are copiously
detailed as regards the mountain Areskuten.

On the death of C. E. Sowerby, the proprietor of the

330 GEOGRAPHICAL BOTANY.

'English Botany/ his successor, J. D. C. Sowerby com-
menced a new series of the parts of this illustrated work,
of which, with the aid of Wilson, Berkeley, Babington,
and Borrer, up to 1844, the first three parts have ap-
peared (Supplement to English Botany, second series,
Nos. 1-3. London). The Botanical Society of London,
following the example of that of Edinburgh, have pub-
lished a catalogue of British plants (The London Catalogue
of British Plants, published under the direction of the
Botanical Society of London. London). Inconsequence
of critical elaboration, this catalogue contains considerably
fewer species (1305 indigenous, and 132 acclimated pha-
nerogamia) than the Edinburgh one, and is ascribed to
the pen of Watson. The ' Phytologist/ a journal which
was noticed in the yearly report for 1842, is still con-
tinued. I may refer to the list of contents given in the
' Botanische Zeitung/

Watson has made some critical remarks upon individual
British plants (London Journal of Botany, iii, pp. 63-81).
Newman has issued a description of British Ferns (A
History of British Ferns and allied Plants. London,
1844). 'The Annals of Natural History' (vol. xiii, xiv)
contain the following contributions to the British Flora :
Ball, on (Enanthe ; Taylor, contributions to our know-
ledge of the Jungermannice ; Harvey, description of the
new Irish genus of Algae, Rhododermis ; Berkeley, con-
tributions to Mycology ; Dickie, critical catalogue of the
Marine Algae existing at Aberdeen ; Spruce, catalogue of
the Mosses and Hepaticae of Teesdale, in Yorkshire ;
Salwey, of the Lichens of Wales ; Graham, on the results
of his journey through Wales ; Babington, on the Irish
Saxifrages.

Babington has shown that Neottia gemmipara Lm.,
the rarest of all the European Orchidaceae, which was
discovered by Drummond near Cork, in 1810, and has
only recently been again found, is identical with the
Spiranthes cernua Rich, of North America (Proceed, of
the Linnaean Society, 1844).

GEOGRAPHICAL BOTANY. 331

Sande Lacoste, and Dozy have been occupied in the
study of the cryptogamic plants of the Netherlands. The
former has made known localities of the Mosses ; the
latter, in conjunction with Molkenboer, has published a
catalogue of the Fungi indigenous to that country, and
some newly- discovered Mosses (both in v. d. Hoeven's
Tidjschrift, f. 1844, p. 165 and 377).

The general works upon the Flora of Germany, men-
tioned in the previous Annual Reports, have been con-
tinued. Four decades of the seventh volume of Reichen-
bach's c Icones/ containing the Aroidese and the allied
groups, have appeared. A cheaper edition, containing
a more copious text, was commenced at the same time,
with the title of ' Deutschland's Flora/ Parts 23 and
24 of the third section of ' Sturm's Flora;' the fifth
volume of Schlechtendal and Schenk's illustrated work ;
and Parts 48-56 of that upon Thiiringia; Parts 34-49
of Link's Publication ; and Parts 2-4 of D. Dietrich's
' Crypt ogamia.'

Rabenhorst has published the first volume of a ' Cryp-
togamic Flora,' containing the Fungi (Deutschland's
Kryptogamen Flora. Bd. i. Leipzig, 1844-8). This
compilation is adapted to the present time, but does not
entirely come up to our expectations. Of the author's
valuable collection of dried Fungi, the seventh, and in
the following year the eighth, " Centurie" have appeared.
Hampe is preparing a similar herbarium of the ' Crypto-
gamia of the North of Germany/ which comprises at pre-
sent 230 Mosses, 80 Hepaticse, and 80 Lichens. (By the
author at Blankenberg, on the Hartz.)

In Wallroth's ' Contributions to Botany/ two parts of
which are before us, individual genera of the flora of
Germany are treated monographically ; especially Agri-
monia, Armeria (with two well-marked Hartz mountain-
plants, Agrim. odorata D. C., Syn. A. procera Wallr.,
and Armeria humilis Lk., Syn. A: fllicaulis Boiss. !
A. Hatteri Wallr.), Lampsana, and Xanthium. Then
follow critical remarks ; as, e. g., upon Senecio paludosus.

332 GEOGRAPHICAL BOTANY.

Salix Uastata, from which Wall roth distinguishes the
form which he discovered on the gypsum-chain of the
southern Hartz mountains, as S. surculosa. Scheele has
continued his work upon German and individual Exotic
Plants, which was noticed in the last Annual Report
(Ratisbon Flora, 1844, and Linnaea, 1844); and Peter-
mann has followed him in attempts of the same kind, to
contribute to our knowledge of native species (Ratisbon
Flora, id.).

Provincial Topographies and Sketches of the Vege-
tation in the Province of the German and Prussian
Flora : Kamp, Catalogue of Plants growing wild around
Memel (Preuss. Provinzialblatter, 1844, p. 451-569);
Leo Meier, On the Flora of Gerdana in Eastern Prussia
(Bot. Zeit., 1844) ; Roeper, Contributions to the Flora
of Mecklenberg (Part 2, Rostock, 1844), representing
the Graminacese in the manner pointed out above ;
Fiedler, Synopsis of the Mosses of Mecklenberg (Schwerin,
1844-8)5 Hacker, Flora of Liibeck (1844-8); K.
Miiller, Contributions to a Cryptogamic Flora of Olden-
burg (Bot. Zeitung, 1844), with additions and correc-
tions by H. Koch (id.) ; Wimmer's Flora of Silesia,
which was mentioned in the Annual Report for 1840,
has appeared in a second and enlarged edition (Breslau,
1844) ; Reichenbach, Upon the Botanical Conditions of
the Flora of Saxony (i. e. Gaea of Saxony, 1843-8),
contains nothing more than a catalogue of rare plants
from the separate districts, in the form of extracts from
the author's 'Flora Saxonica;' Pfeiffer, Sketch of the
Plants hitherto found in Kur-Hesse (Cassel, 1844-8);
this is to be regarded as preliminary to a critical Flora of
Hesse, and contains a large number of new localities,
especially on the basaltic mountains of Cassel ; by the
same author, A few words upon the Subalpine Flora of
Meissner (loc. cit., 1844); Wirtgen, Supplements to the
Flora of the Prussia Rhine Provinces (Verhandlungen des
naturhistorischen Vereins der Preussischen Rheinlande,
Jahrg. 1) ; Thieme, Catalogue of the Plants growing at

GEOGRAPHICAL BOTANY. 333

Hainsberg, in the Territory of Aix (Katisbon Flora,
1844, p. 209-21); Lohr, Manual of the Mora of
Treves and Luxembourg, with a notice of the surrounding
districts (Treves, 1844-8); Lechler, Supplement to the
Flora of Wiirtemberg (Stuttgard, 1844-8) ; Sailer,
Flora of Linz (Linz, 1844-8), an extract from the Flora
of Upper Austria, mentioned in the Annual Report for
1841 ; Sauter, Report upon a Journey to Luiigau (Ra-
tisbon Flora, 1844, p. 813-16).

E. v. Berg, at Lauterberg, on the Hartz mountains,
endeavoured to prove that the Coniferse are gradually be-
coming more widely distributed in the north of Germany
(Das Verdrangen der Laubwalder durch die Fichte und
Kiefer. Darmstadt, 1844-8). The fact, in the case of
the Hartz mountains, rests upon authentic testimony ;
but how far this change, which in many places has been
completed in the space of twenty years, has been pro-
duced by external natural conditions, or merely by the
economic management of the forests, is difficult to ascer-
tain. In Liineburg also, where, e. g., in the struggle
between the two methods of culture, it was not decided
in favour of the pine until after the lapse of a century,
as also in Soiling, on the Upper Weser, where deciduous
forests are still very extensive, the same conditions have
prevailed as on the upper Hartz mountains. On the
western Hartz mountains, the red pine generally succeeds
the beech ; but in some parts, on the removal of the
latter, the remains of oaks have been found as high as a
level of 2000', i. e. an elevation at which they have long
since ceased to grow. When we consider that the tree-
limits on the Hartz mountains lie extremely low, in
comparison with those of the north of Europe, and that
even the Coniferse do not ascend higher on the Brocken
mountains than at 9 10 further north in Norway, the
fact of the culture of the oak and beech at a former period,
would render it, at any rate, tolerably probable that
secular changes had taken place in the climate, by means
of which the distribution of the forest trees had been

334 GEOGRAPHICAL BOTANY.

produced, and by which Steenstrup's succession of forest
growth in maritime countries would be brought into
connexion with the extermination of the Coniferse in the
elevated regions of the upper Hartz mountains.

The work of Fuchs, on the Venetian Alps, contains
an account of the limits of vegetation in the southern
dolomitic Alps, especially the district of Agordo ; it fills
up an important gap in the observations upon the ver-
tical distribution of the Alpine plants (Vienna, 1844, fol.)
Unfortunately, however, in the case of most of the plants,
the lower limits of altitude only are given ; and of these,
a local value only can be attributed to many measure-
ments. The results, expressed in French feet, are as
follows :

a. Upper limits.

Ficus Carica, and limit of the cultivation of the vine, 1500'. (At Agordo,
Vitis grows very luxuriantly at a level of 2000', but no wine is made.

Castanea vesca, 2000' at Agordo.

Juglans regia, 3500' at Frassene.

Zea Mays, 2500' in the valley of Cordevolethal.

Cerealia, excluding wheat, 4400' at the Col di S. Lucia; 4600' at
Buchenstein.

Dense Forest of Coniferee, 5500'. In the regions of the mountain-pine,
individual larches and fig-trees ; 6309' at Sasso di Palma.

Fagus sylvatica, 5000' ; e. g. at Monte Luna, 4915', still higher at Bosco
Medona and in the Val Pegolera.

Pinus Cembra, 6665' at the Col di Lana.

Upper limit of the Phanerogamia, = 9000' ; Aretia Vitaliana, and some
Saxifrages.

b. Lower limits.

^Ranunculus aconitifolius, 3500'. Hutchinsia rotundifolia, 7000'.

montanus, 7000'. Papaver pyrenaicum, 5500'.

glacialis, 8000'. Viola biflora, 3500'.

Pyrenaicus, 8000'. Silene acaulis, 5500'.
Anemone baldensis, 4500'. jmmilio, 7000'.

Aconitnm Anthora, 4500'. Cerastium latifolinm, 6500'.

Napellus, 6500'. Cytisus alpinus, 1300'.
Stoerkiannm, 6500'. purpurens, 2000'.

Arabis cterulea, 7000'. Trifolium alpinum, 5500'.

Hutchinsia alpina, 7000'. Phaca astragalina, 6500'.

GEOGRAPHICAL BOTANY.

335

Phetca alpina, 6500'.
Hedysarnm obscurum, 7000'.
Dryas octopetala, 2000'.
Potent ilia caulescens, 1300'.

nitida, 6500'.
Geum montanum, 5500'.

reptans, 8000'.
Sibbaldia procumbens, 5500'.
Rosa alpina, 5500'.
Sedum atratum, 7000'.
Rhodiola rosea, 7000'.
Saxifraga Aizoon, 1300'.

aizoides, 1500'.

casia, 1500'.

rotundifolia, 2000'.

mutata, 2500'.

Burseriana, 2500'.

cuneifolia, 3500'.

stettaris, 5500'.

aspera, 5500'.

controversa, 6500'.

muscoides, 6500'.

planifolia, 7000'.

androsacea, 7000'.

sedoides, 7000'.

bryoides, 7000.

oppositifolia, 8000'.
Bupleurum graminifolium, 6500'.
Lonicera nigra, 4500'.

'alpigena? 4500'.
Valeriana saxatilis, 1300'.
^5^r a/^m^ 3 1500'.
Tussilago alpina, 2000'.
Cacalia alpina, 4500'.
Arnica montana, 2000'.

Bellidiastrum, 1300'.
Gnaplialium Leontopodium, 1500'.
Chrysanthemum alpinum, 7000'.
Anthemis alpina, 6500'.
Achillaea Clavennae, 4500'.
moschata, 7000'.
Doronicum scorpioides, 7000'.
Aronicum Clusii, 7000'.

Senecio abrotatiifolius, 5500'.

carniolicus, 7000'.
Cirsium, ochroleucum, 2500'.

spinosissimum, 5500'.
Carduus defloratus, 5500'.
Saussurea alpina, 7000'.
Sonchus alpinus, 4500'.

# comosum, 1300'.
Scheuchzeri, 1300'.
hemisph&ricum,, 5500'.
orbiculare, 550Q'.
Sieberi, 7000'.
pauciflorum, 7000'.
barbata. 4500'.
Morettiana, 4500'.
Rhododendron hirsutum, 1300'.

Chamacistm, 1300'
Arbutus uva ursi, 2500'.
/j9^, 5500'.
Azalea procumbens, 7000'.
Vacdnium Myrtillus, 2000'.

r#w *^tf, 2000'.

Primula Allionii, 2500'.
glutinosa, 7000'.
minima, 7000'.
longiftora, 6500'.
Auricula, 6500'.
Soldanella alpina, 2500'.
minima, 2500'.
CWaw* Matthioli, 7000'.
Androsace alpina, 7000'.

obtusifolia, 7000'.
Jr^w Titaliana, 8000'.
Pinguicula alpina, 2000'.

grandiflora, 2000'.
Gentiana acaulis, 1300'.

germanica, 1300'.
utriculosa, 2000'.
cruciata, 3500'.
asclepiadea, 3500'.
7wfo, 3500'.
punctata, 5500'.
bavarica, 5500'.

336 GEOGRAPHICAL BOTANY.

Qentiana nivalis, 5500'. Betonica Alopecuros, 1300'.

pumila, 5500'. Myosotis nana, 8000'.

Linaria alpina, 1300'. Globularia nudicaulis, 1300'.
Euphrasia tricuspidata, 1300'. cordifolia, 1300'.

Salisburgensis, 1500'. Daphne striata, 1500'.

Pedicularis tuberosa, 4500'. Pinus Pumilio, 1400'. Between
rostrata, 6500'. Agordo andPeron.

verticillata, 6500'. Nigritella angustifolia, 4500'.

rosea, 6500'. Himantoglossum viride, 4500'.

Bartsia alpina, 6500'. Crocus vernm, 2000'.

Paderota Bonarota, 1500'. Czackia Liliastrum, 2500'.

Veronica alpina, 4500'. Luzula nivea, 5500'.

aphylla, 4500'. Carex atrata, 5500'.
Horminum pyrenaicum, 1300'. firma, 5500'.

Giacich has enumerated the rare plants of Monte
Maggiore, in Istria (Ratisbon Flora, 1844, pp. 274-6).
Hauffel gives a sketch of the Carices of Hungary,
Croatia, Slavonia, and Siebenbiirgen (Id. pp. 527-36).
The author here again refers his C. rhynchocarpa to
C. brevicollis Lam., and regards C. saxatilis Baumg. as
C. dacica.

Moritzi has written a new Manual of the Swiss Flora
(Die Flora der Schweiz. Zurich, 1844-8). Trog has
published a Catalogue of Swiss Fungi (Berner Mitthei-
lungen, pp. 17-92), in which 1121 species are mentioned.

The upper limit at which the larch occurs on the south
side of the Mont Blanc chain at Cramont, in Courmayeur,
was found by the measurement of Forbes, to be 7200'
Engl. ; and on the north side, on the rocks les Echellets
which belong to the Mer de Glace, 6800'. (Travels
through the Alps of Savoy. Edinb., 1843, pp. 68 and
215.)

The seventh and eighth centuries of F. Schultz's 'Flora
Galliae et Germanise exsiccata' have been issued, and are
accompanied by critical remarks upon individual plants
(see Bot. Zeitung, 1845). By the same author, four
French plants are proposed as new in the Ratisbon Flora'
(1844, pp. 806-9); Orobanche brachysepala Sch. accord-
ing to the description given, and comparison with the

GEOGRAPHICAL BOTANY. 337

original plants, is identical with 0. apiculata Wallr.
Rchb. (Spicil. rum., 2, p. 58) ; 0. macrosepala is pro-
bably my 0. Bartlingii, a name which obtained priority
by several months.

French local Floras : J. Lloyd, ' Flora de la Loire in-
ferieure' (Nantes, 1844, 12); Guepin, 'Supplement a la
Flore de Maine et Loire' (Angers, 1842).

Martins has worked out an exposition of the climatal
contrasts which occur within the boundaries of France
(les Regions Climatoriales de la France), in the ' Biblio-
theque de Geneve, 1844, pp. 138-60, and pp. 347-50.
The author distinguishes the five following climates in
France.

1 . Climate of the Vosges. This comprises a district in
the north-east of France, which is bounded by the citiss
of Basle, Dijon, Auxerre, and Mezieres. Mean tempera-
ture = 9 6C. The relatively most intense winters pre-
dominate in this region, the difference between the mean
summer and winter heat amounts to 18 C. (18 6 and
6 C.) ; the greatest cold observed in Strasburg and
Metz amounted to about 23 C. The mean quantity of
rain (from meteorological observations made at Strasburg,
Muhlhausen, Nancy, Metz, and Geneva) =669 mm. ;
of this, 19 p. c. fall in the winter, 23 in the spring, 31
in the summer, and 27 in the autumn. Average num-
ber of rainy days = 137. Predominant winds, those
from the south-west and north-east.

2. Climate of the Seine , or north-west of France, as
far as the Loire and Cher. Mean temperature 10 9 C.
Difference between the mean summer- and winter-tem-
perature = 13 60.; diminishing in the direction from
Brussels (=14 3 C.) to Brest (= 10 8 C.) ; the former
average value is the arithmetical mean of observations
made at Dunkirk, Arras, Abbeville, Paris, Cherbourg,
Angers, and Denainvilliers. Average amount of rain
= 548mm.; in Finisterre, however, it amounts to 900mm.
(from observations made at Paris, Brussels, and Denain-
villiers, 21 p. c. of the rain falls in winter, 22 in spring, 30

338 GEOGRAPHICAL BOTANY.

in summer, and 27 in the autumn. Average number of
rainy days =140. The prevailing wind is the south-
west, the next is the north-east.

3. Climate of Garonne, or south-west of France^ as
far as the Pyrenees. The eastern boundary is situated in
Auvergne, but cannot at present be accurately defined ;
it probably includes the plateau of Auvergne, and follows
the course of the Rhone and Saone. Mean temperature
=: 12 7 C. Difference between summer- and winter-
heat =: 160 C. ; on account of the smaller extent of the
coast-line, the marine climate is less developed here than
in the north-west ; mean summer- temperature = 20 6 C.,
winter-temperature = 5 C. Greatest intensity of cold .at
Poitiers, La Rochelle, Toulouse, and Agen, where it
attains to 1 2 C. Average amount of rain =586 mm.;
of which, 25 p. c. fall in winter, 21 in spring, 23 in
summer, and 34 in autumn. Average number of rainy
days = 130. The prevailing wind is thje south-west,
which in the neighbourhood of the Pyrenees passes into
the west.

4. Climate of the Rhone, comprises the valley of the
Rhone, from Dijon and Besancon to Viviers, and the
mountainous regions of the Higher Alps ; the boundary
in the department of the Lower Alps is at present unde-
fined. Mean temperature = 11C. Difference between
summer and winter = 18 6 C. Mean temperature of
summer = 21 3 C., of winter 2 5 C. Average quantity
of rain = 946 mm., i. e. the greatest amount precipitated
throughout the whole of France ; of which 20 p. c. falls
in the winter, 24 in the spring, 23 in the summer, and
34 in the autumn. Number of rainy days in the valley
of the Saone = 120-30, in the valley of the Rhone =
100-15. Prevailing winds, north and south.

5. Mediterranean climate. The northern boundary
runs through the Rhone at Viviers, near Montelimart,
thence follows a line drawn on one side to Montpellier,
on the other to Marseilles, and, lastly, comprises the coast-
districts of Provence and the regions of Aude as far as

GEOGRAPHICAL BOTANY. 339

the Pyrenees. Mean temperature =14 8 C. Mean
summer temperature = 22 6 C., and winter temperature
= 6 5C. Greatest intensity of cold observed 1 1 5 C.
Average amount of rain = 651 mm. ; of which 25 p. c.
fall in the winter, 24 in the spring, 11 in the summer,
and 41 in the autumn. Prevailing wind, north- west.
(Mistral.)

A work by Grenier, relating to the botanical condi-
tions of the French Jura, appears of importance ; at
present, however, I am only acquainted with it from Von
Schlechtendal's review (These de Geographic Botanique
du Dep. de Doubs, Strasbourg, 1844-8). According to
this work, the upper limit of the oak here occurs at an
altitude of 6-700 metres, that of the beech at 8-900
metres ; above these deciduous trees comes the Coniferous
region, covered with both kinds of fir-trees.

Lloyd's Flora of the Mouth of the Loire also notices
the local conditions of vegetation. The diffusion of several
plants belonging to the south of Europe, along the sea-
beach, as far as the 47th degree of latitude, is charac-
teristic : e. g. on the lagunes, Inula crithmoides, Sonchus
maritimus, several Statices, Salicornia fruticosa, Scirpus
Savii y Spartina strict a ; on the downs, Matthiola sinuata,
Silene portensis, Tribulus terrestris, Otanthus maritimus,
Ephedra distachya, Pancratium maritimum, &c. But on
the heaths of Bretagne are also found Erica ciliaris,
vayans, and scop aria, Simethis bicolor Kth. (Phalangium
D. C.), Aspkodelus albus, Pingmcula Lusitanica., Serapias
triloba, in conjunction with northern plants, as Ulex
Europ&us, Narthecium ossifrayum, Anagattis tenella, Hy-
pericum elodes, Myrica Gale, and Alisma ranuncidoides.

To this place belong, on the French coast of the
Mediterranean, the investigations of Duchartre upon the
vegetation of the district around Beziers in the Dep.
Herault (Comptes rendus, 1844, v, 18, pp. 254-9). This
work gives an accurate and complete survey of the vege-
table formations which occur there. The author divides

340 GEOGRAPHICAL BOTANY.

them into two principal classes, according as their growth
is consequent on proximity of the sea or not.

1. The following formations belong to the coast-
plants : a. Formation of the Dunes. Herbs or low shrubs,
which are either of very pubescent or of glaucescent
tint. To the former belong, e. g. Mattkiola sinuata,
Medicago marina, Orlaya maritima, Mercurialis tomen-
tosa. Diotis candidissima ; to the latter, Eryngium mari-
timum, Echinophora, Euphorbia Par alias, and Crucianella
maritima. The shrubby plants consist of Astragalus
massiliensis and Ephedra distacliya. As regards the
number of forms, the Grasses predominate (12 species
are known), the Cruciferae come next, with 4 species, the
Leguminosae and Euphorbiaceae number 3 species, and
the Chenopodeaceae, Polygonaceae, and Synantheraceae :
altogether more than 40 species grow there. Among the
dunes 2 allied species of Juncus are found (J. acutus and
maritimus), a formation peculiar to the humid soil, which
in the Landes is denominated Joncasses, and forms the
transition to the following formation.

b. Formation of the Salt-water Marshes. Shrubs and
herbaceous perennials, with succulent leaves. Cheno-
podeae and Statices predominate here, both as regards
the number of individuals and of species : among woody
plants, Tamarix Gallica is found arborescent. Charac-
teristic forms among the Chenopodeaceae (11 sp.) : Che-
nopodium fruticosum, Ch. setigerum, Salicornia, 3 sp.,
Salsola, 2 sp., Atriplex, 3 sp. ; of the Statices (5 and
more species) St. oleifolia, bettidifolia, and ferulacea ;
of other plants (15 sp.), Frankenia, 2 sp., Spergularia,
2 sp., and Artemisia Gallica. The Graminaceae are here
represented by Crypsis schcenoides only.

2. The plants independent of the influence of the sea,
resolve themselves into formations of a moist and dry
soil ; the latter are either independent of the cultivation
of the land or not so.

A. Water-plants.

GEOGRAPHICAL BOTANY. 341

a. Fresh-water Formation. Amongst numerous Gra-
minaceae, Cyperaceae and Naiadeae, with Nymphaeaceae
and Typheae, it possesses but few forms which are charac-
teristic of the climate ; as, e. g. Vallisneria spiralis and
Marsileapubescens, Ten. (M. Fabri, Dun.)

b. Formation of those soils which are occasionally
overflowed, are characterised, e. g. by Mentha cervina.
These are the localities of Cicendia Candollei and Conyza
sicula.

B. Plants of the uncultivated part of the district. The
author believes that three or four formations may be dis-
tinguished, of which the first, that of the Cistaceae, is
more distinct than the rest are from each other.

a. Garrigues, i. e. Formation of the Cisti. A stony
soil is densely covered with shrubs of Cistus or some
other firmly interlaced and frequently thorny bush. These
shrubby forms are the following : Cistus crispus, salvi-
folius, albidus, and monspeliensis ; Ulex provincialis and
Europaus ; Daphne Gnidium ; Quercus coccifera ; Erica
scoparia and cinerea, Calluna vulgaris ; PUillyrea angus-
tifolia and latifolia ; Lavandula Stcechas ; Osiris alba ;
Junipcrus Oxycedrus and communis, and Rosmarinus offi-
cinalis. Among other plants, the following are charac-
teristic : a number of species of Helianthemum, growing
with the Cisti, some Euphorbias, Santolina, Helichrysum
Stcechas, Aphyllanthes, &c. From this formation also
more than 40 species are enumerated.

d. Duchartre has not been able to distinguish the pecu-
liarities of those surfaces and hills (campi) which are
covered with shrubs and annuals, and to illustrate them
according to their characteristic vegetable forms. We
shall, therefore, pass over this part of his memoir, and
merely mention some of the more rare species which
belong here : Biscutella coronopifolia, Linum salsoloides,
Centaurea Pouzini, and Echimn Pyrenaicum.

c. Plants of the cultivated soil.

a. Formation of the ruderal plants. The species are
all widely diffused.

342 GEOGRAPHICAL BOTANY.

b. Plants accompanying those which are cultivated.
The author makes several divisions of these, which it is not
necessary to detail. The number of species enumerated
is very considerable, but they are not characteristic of the
south of France, as distinguished from other countries on
the Mediterranean.

c. Formation of the meadow-lands. The same remark
applies to this : Euphorbia pilosa and Iris spuria, how-
ever, deserve to be mentioned.

d. Formation of the forests. The evergreen forests
consist of Quercus Ilex : there are no others. Under-
wood : Pis facia Lentiscus and Terebinthus, Erica arbor ea
and Calluna, ISarothamnus scoparius, Cytisus capitatus,
Genista Scorpius, Spartium junceum, &c.

From the appended sketch of the cultivated plants, it
is seen that the preparation of soda from Halophytes has
entirely ceased in that district, that the cultivation of
the olive is very much on the decrease, in consequence of
several cold winters having destroyed the plantations, and
that latterly attempts have been made to cultivate Eicinus
on a large scale. The principal production of Beziers is
wine ; the cerealia do not suffice for home consumption.

Desrnoulins has given a description of his botanical
journey in the Pyrenees, during which he made some
observations upon the vertical limits of the Alpine flora
of the Pic du Midi (Etat de la Vegetation sur le Pic du
Midi de Bigorre. Bordeaux, 1844, 8vo.) We extract
from it the following additions to the earlier statements of
De Candolle and Ramond :

Cochlearia pyrenaica, 5500' 6000'.
Herniaria pyrenaica, 3000' 7500'.
ParonycMa polygonifolia, 6000' 7500'.
serpyllifolia, 7500 8400'.
Astragalus depressus, 6000' 7500'.
Vicia pyrenaica, 8500'.
Carduus carlinoides, 6000' 8100'.

carlinifolius, 3000' 6900'.
Cirsium eriopJiornm, 0' 6600'.
Scabiosa pyrenaica, 8400'.

GEOGRAPHICAL BOTANY. 343

Pedicularis pyretiaica, 9000'.
Crocus nudiflorus, 7500'.
Anictangium cilia turn, 8400'.
Pannelia chrysoleuca, 5400' 9000'.

cartilaginea, elegans, cinerea, badia, 9000'.
Lecidea vesicularis biformis, 6000' 7500'.

polycarpa, atrobrunnea, morio, geogmphica, umbilicata, 9000'.
Umbilicaria cylindrica, 6000'--9000'.

Some interesting letters, written by M. Willkomrn
during a journey in Spain, have been published in the ' Bo-
tanische Zeitung' (1844-5). They commence in May 1844,
bearing date from Valencia, where the author remained
until the middle of June. He then went to Madrid,
botanised at Aranjuez in the beginning of July, passed
over the Sierra Morena, reached Granada, and during
the latter part of the summer and the autumn, explored
the Sierra Nevada and the Alpuxarras. In the present
report, we shall confine ourselves to the first part of the
journey, intending to recur to the notices regarding the
south of Spain next year, when the conclusion of Boissier's
illustrated work, together with Willkomm's observations
of 1845, will conjointly furnish a more copious illustra-
tion of that part of the subject. In the Huerta of Va-
lencia, the original vegetation is greatly displaced by
cultivation : wheat, rice, and hemp are principally culti-
vated ; mulberry trees, olives, and fruits belonging to the
south are common,, date-palms, from 40 to 60 feet
high, are frequently met with. On the Lagune Albufera
is a wood of Pinus ffalepensis, containing abundance of
the original plants of this district : the underwood here
consists of Quercus coccifera, Myrtus, and Cham&rops,
and growing with them we find Pistacia Lentiscus,
Rhamnus lycioides, Erica arborea, Eosmarinus, Juniperus
Oxycedrus, and Euscm aculeatus. The adjacent sandy
hills contain Cistus albidus and salvifolius ; Passerina
hirsuta and Solatium sodomeum, the stems of which are
of the thickness of the arm.

The Sierra de Chiva, 1 2 miles north of Valencia, belongs
to the limestone mountains, which separating from the

344 GEOGRAPHICAL BOTANY.

Spanish plateau, between the Ebro and Xucar, traverse
the province from west to east as far as the sea. This
broad mountain -range, which is about 6000' in height,
and intersected with deep Barrancos, was once covered
with forests of Coniferse, the only remains of which at
the present time are isolated stems of Pinus Halepensis.
The dry slopes, which are almost entirely free from springs,
are now overgrown with a low bush (Montebaxo), the
extreme summits only being bare. Willkomm admits
the following stages in the Mediterranean vegetation of
this region, which attains an unusual elevation, ascending
to 4000'.

500'. To about this height the Opuntias and
Agaves extend, together with the culture of Ceratonia.
The Montebaxo consists of Chamber ops, Erica arborea,
Daphne Gnidium, Retama sph(erocarpa, Ulex, Rosmarinus,
and some oaks.

500' 2000', i. e. as far as the upper limits of Cha-
mcerops (also of Retama, Juniperus Oxycedrus, and Pis-
tacia Lentiscus). Rosmarinus and Chamarops predomi-
nate; in addition to those already mentioned, Erica
arborea from among those of the first stage, and Rhamnus
lycioides, Pistachio, Terebinthus, and some Cisti are here
first met with. Characteristic Grasses : Macrochloa te-
nacissima and Stipa juncea.

2000' 4000' up to the limits of the cultivation of the
olive and wheat. The greater part, however, of the slopes
at this level consists of uncultivated mountain-land. In
a Montebaxo, the principal plants associated here with
Rhamnus, Rosmarinus, Erica, and Cisti, are Juniperus
Phcenicea, Fraxinus sp., Arbutus unedo, and Quercus Ilex.

Isolated pine-trees and a Montebaxo formed of Vlex
Australis and Juniperus Phcenicea characterise the re-
gion extending from 4000' 5500', which may be dis-
tinguished from the Mediterranean by the occurrence of
the plants of the north of Europe. On the summit of the
Monte de la S. Maria (5500' 6000'), of woody plants,
Arctostaphylos uva ursi, Taxus, and some Cotoneasters

GEOGRAPHICAL BOTANY. 345

are also found ; and with them few shrubs only and a
single species of Saxifrage.

V. Martens, in a general work, has described the bota-
nical geography of Italy from literary sources (Italy,
Stuttgard, 1844, 8vo, 3 vols.)

Works upon the Flora of Italy. The first two parts of
the sixth vol. of Bertolini's Flora Italica, which treat of
the 14th class, have appeared (Bologna, 8vo.) The Flora
of Nice, by A. Risso (Nice, 1844, 8vo), is of no scientific
value. We have not yet received Cesati's paper upon
that of Lombardy (Saggio sulla Geographica botanica e
sulla Flora della Lombardia. Milano, 1844, 8vo, p. 74).
Purcinelli Additamentum ad synopsin plantarum in agro
Luccensi sponte nascentium (in the Giornale Botanico
Italiano. 1844, pp. 118-123). Savi Florula Gorgonica
(id. pp. 243-283), a catalogue enumerating 290 sp. of
vascular plants observed in Gorgona, a small island oppo-
site Leghorn, and covered with Cisti, Ericas, and Legu-
minous shrubs, may be considered as a companion to the
Flora of Capraja, published some years ago by Moris
and Notaris. De Notaris, Appendix to his Specimen
Algologise Ligusticse (id. pp. 191, 311). Meneghini,
Algarum species nova3 vel minus nota3 (id. pp. 296-306),
33 species from the coasts of Italy and Dalmatia. The
fourth part of Alghe Italiane e Dalmatiche, by the same
author, has appeared (Padova, 1843, 8vo). Ten ore has
shown that the Dalmatian Arenaria Arduini is identical
with his former A. Rosani (Ren die. Acad. 1842, p. 266).
V. Heldreich describes four new Sicilian plants (Ratisbon
Flora, 1844, p. 65): 1 Heliantkemum, 1 Elichrysum, 1 Cen-
taurea, and 1 Lit/iospermum. Nyman's Observations in
Floram Siculam (Linna3a, 1844, pp. 625-665) contain a
catalogue of his collection which is for sale in Sweden,
with descriptive remarks. The only new plant is Parie-
taria populifolia, N. from Malta.

Link distinguishes a new Erica ant hum, obtained from
Spalatro (Sitz. des Ges. naturf. Freunde, 1844, in the
Ratisbon Flora, 1845). Visiani raises Turinea Neu-

346 GEOGRAPHICAL BOTANY.

mayerina Vis., which was figured, in the Flora Dalma-
tica, into a separate genus as Amphoricarpos (Giorn. Bot.
It,, vol. i, p. 196).

Ebel's essay on Montenegro (Zwolf Tage auf Monte-
negro, Hft. 2. Konigsberg, 1844, 8vo), contains a cata-
logue of all the Phanerogamous plants hitherto observed
in Dalmatia (2003 sp.), with a statement of the frequency
of their occurrence, expressed in a manner peculiar to
the author, but the localities are not given. It contains
preliminary observations upon the statistical relations of
the flora of Dalmatia, in which the most abundant families
form the following series, according to the number of
species contained in them : Synantheraceae (225 sp.),
Leguminosae (220 sp.), Graminaceae (142 sp.), Cruciferae
(107 sp.), Umbelliferae (103 sp.), Labiatae (91 sp.), Ga-
ry ophyllacece (85 sp.), Scrophulariaceae (82 sp.), Liliaceae
(61 sp.), Rosaceae (59 sp.), Ranunculaceae (54 sp.), Orchi-
daceae (46 sp.), Cyperaceae (43 sp.), Boraginaceae (42 sp.)
The reports upon the vegetation of Montenegro itself, the
productions of which, according to the author, entirely
agree with those of Dalmatia, belong here. This small
tract of land, which is covered with arid, rocky mountain-
pastures, and elevated into limestone summits, which are
either barren or slightly surrounded with forests of pines,
and from which narrow fluviatile valleys descend to the
sea of Scutari, is extremely unfruitful from a deficiency
of soil and water. Nevertheless, the plants appear, as in
Dalmatia, to be various, 450 sp. having already been
mentioned by the author : there are no new ones among
them, the two which are proposed as new are untenable.

In my work upon Rumelia and Bithynia (Spicilegium
Floras Rumelicae et Bithynicae, exhibens synopsin plan-
tarum, quas anno 1839 legi: accedunt species, quas in
iisdem terris lectas communicarunt Friedrichsthal, Fri-
waldzki, Pestalozza vel plene descriptas reliquerunt Bux-
baum, Forskal, Sibthorp, alii; vols. i, ii, Brunsnigae,
1843-4, 8vo), 2300 Phanerogamous plants are treated
of systematically, and in regard to their geographical dis-

GEOGRAPHICAL BOTANY. 347

tribution. The families containing most species form
the following series : Synantheracea3 (264 sp.), Legumi-
nosae (203 sp.), Graminaceae (156 sp.), Labiatae (134 sp.),
Caryophyllaceae (130 sp.), Cruciferae (121 sp.), Umbelli-
ferae (114 sp.), Scrophulariaceae (90 sp.), Ranunculaceae
(78 sp.), Rosaceae (68 sp.), Boraginaceae (55 sp.), Lilia-
ceae (53 sp.), Rubiaceae (48 sp.), Campanulaceae (41 sp.),
Orchidaceae (41 sp.), Cyperaceae (41 sp.) When this
series is compared with that given above for Dalmatia,
the increase in the Labiatae and Caryophyllaceae becomes
one of the characteristic peculiarities of Rumelia. The
former family does not reach the centre of its distribution
through the south of Europe until we arrive at Greece ;
but the Silenaceae, which abound in endemic forms of
Dianthus and Silene, do not appear to be anywhere more
numerous than in Rumelia. The increase in the Ranun-
culaceae, Boraginaceae, and Campanulaceae is also worthy
of consideration ; but I must confine the deductions to these
few facts, since if carried further than the extent of our
present knowledge admits, they would lose in truth.
The extent of our knowledge of the flora of Rumelia is
much better shown by the examination of those vegetable
forms which are endemic to that country, than by sketches
of the entire vegetation, in which so many constituents
are still wanting. Of these 2300 species of plants, about
the seventh part are peculiar to the peninsula of Europe :
from these about 80, which have only been found in
Bithynia, are excluded, a great part of which, however,
will probably be found also on this side of the Bosphorus.
Moreover, if we take into account the distribution of
Greek plants over the south, and of Dalmatian over the
west of Rumelia, we may consider more than two thirds
of the endemic plants of the south-east of Europe as
known. Summary of the endemic plants of Rumelia :
23 Leguminosae, principally species of Trifolium (5), and
Astragalus (9), mostly belonging to the evergreen region ;
5 Rosaceae, of these, 3 Dryadeae to the mountainous re-
gion ; 2 Rutaceae (Hoplophyllum) ; 4 Euphorbias, of which

348 GEOGRAPHICAL BOTANY.

2 belong to the alpine region ; 2 Geraniaceae to the alpine
region; 25 Caryophyllaceae, especially species of Silene
(6), and Dianthus (10), only 5 Alsineae : species from all
three regions, but the pinks mostly indigenous to ^the
central European and alpine ; 5 Hypericineae (Hypericum}
from the evergreen region ; 14 Cruciferae, one half of
which consist of alpine species of Arabis, Cardamine,
Koniga, Thiaspi, and Eunomia ; 1 5 Ranunculaceae, with
7 species of Rammculus, mostly from the evergreen region ;
2 Crassulacese ; 3 Saxifrages from the alpine region ; 21
Umbelliferae, increasing towards the coast ; 2 Ericaceae :
Erica verticillata and Arbutus Andrachnm ; 3 Primulaceae ;
26 Scrophulariaceae, principally alpine Pedicular es (3),
species of Veronica (4), Digitalis (3), Scropkularia (4),
and Verbasca of the evergreen region (8) ; 2 Orobanches ;
9 Boraginaceae, among these 4 species of Alkanna, 2 of
Borago ; 20 Labiatae, of these 6 species of Stachys in
both the lower regions ; 9 Rubiacea3 in the evergreen and
alpine region (instead of the term Galium trie/top /iorum,
which was elsewhere proposed at the same time, I prefer
that of G. Trichodes) ; 2 Valerian aceae ; 9 Dipsaceae ; 40
Synantheraceaa, principally Anthemideae and Cynareae,
mostly from the genera Anthemis (6 : mostly in the ever-
green region), Achillea (5 : mostly in the alpine region),
Senecio (4), Centaurea (5), Cirsium (5); 13 Cainpanula-
ceae, of which ] were Campanula, most of which belonged
to the evergreen region ; 2 Amentaceae : Quercus ^Eyilops
and infectoria ; 3 Coniferae : Pinus maritima in the lower,
Juniperus sabinioides in the middle, and Pinus Pence on
the boundary of the alpine region ; 3 Orchidaceae ; 4 Iri-
daceae, species of Crocus in the evergreen region; 12
Liliaceae, e. g. Ornithogalum (3) ; 2 Cyperaceae ; 1 1 Gra-
minaceae from all three regions. The remaining endemic
plants are as yet single members of their families : the
Bithynian, &c. Of the Cryptogamia, we are not yet
acquainted with 200 species.

Heldreich observed at Athens a form of Arbutus, which
was probably^. Uybrida Ker., but is regarded by him as

GEOGRAPHICAL BOTANY. 349

a distinct species, intermediate between A. Unedo and
Andrachne (Ratisbon Flora, 1844, p. 13). He denies
its hybrid origin, because A. Unedo flowers in October
and November, A. Andrachne in February and March ;
I have, however, met with both plants in flower at the
same time in Bythynia.

II. ASIA.

Among the endemic plants of Bithynia described in
the 'Spicileg. Rumelic./ part of which belong to the
evergreen coast-region, part to the high mountains of
Olympus and Bolu, the following are the principal fami-
lies represented : 5 Leguminosse (mostly Trifolia) 2
Geraniaceae ; 5 Caryophyllacese (consisting of 3 Silenese
and 2 Dianthi, all from Olympus) ; 4 Hyperica, 9 Cruci-
ferse (all from Olympus, and consisting of 3 sp. of
Arabis, 2 sp. Eunomia, &c.) ; 3 Papaveracese ; 2 Ranun-
culacege ; 5 Umbelliferse (mostly from Olympus) ; 4
Scrophulariaceae; 2 Boraginacese; 3 Labiatse; 3 Rubiaceae;
13 Synantheraceae ; 4 Campanulaceae ; 3 Liliaceae; 3
Graminaceas, &c.

The oriental Umbelliferae have been worked out by
Boissier (Ann. Sc. Nat. 1844), comprising 300 species.
The number of species proposed as new is very large.
The new genera distinguished are the following: Lereschia
(Cryptotcenia Thomasii D. C.) ; Elwendia, from Persia,
near Carum ; Microsciadium (Cyminum minutum Urv.) ;
Muretia (Bunium sect. Cliryseis D. C.) ; Diplotania from
Persia, near Peucedanum ; Stenotania from the same
place, near Paslinaca; Ducrosia (Zozimice, sp. D. C.) ;
Ainsworthia (Hasselquistia cordata L.) ; Trigonosciadium
from Mesopotamia, near Heradeum ; Synekosciadium
(Heracl. Carmeli Lab.) ; Polylophium, (Thapsiese) from
Persia ; Smyrniopsis near Smyrnium ; Meliocarpus near
Prangos; Turgeniopsis (Turgenia foenicutacealfzl.); Liscea

350 GEOGRAPHICAL BOTANY.

(Turgenice, sp. D. C.) ; Rhabdosciadium, one of the Scan-
dicinese from Persia ; T/iecocarpus, from the same place
and from the same division ; Osmosciadium, one of the
Coriandrese, from Cappadocia.

C. Koch's travels to the Caucasus (Reise durch Russ-
land nach dem Kaukasischen Isthmus in den J. 1836-8.
Bd. i, ii. Stuttgard, 1842-3) contains reports upon the
autumnal vegetation of Ossetia and Imiretia, as also upon
the vernal flora of Russian Armenia ; the author's inves-
tigations were subsequently interrupted by protracted
illness, but he finally resumed them in a second journey.
In the military road of the Caucasus, Koch represents
the prairies of Kabarda, near Uruch, as very luxuriant
and abundant in plants ; herbs and the Grasses grow here
in such luxuriance, that a man can readily conceal him-
self without lying down (i, p. 250). The Graminaceae
are mostly the same as the meadow-grasses of central
Europe, whilst among the shrubs many Caucasian species
are met with ; they are diffused by the rivers over these
surfaces which are situated opposite to the high mountain
chain. By this circumstance and the development of the
vegetation in the height of summer, when the Russian
heaths are burnt up, the meadows of Kabarda differ
essentially from the steppes, with which C. Koch has
compared them. In fact, judging from certain kinds of
plants, the steppe climate still prevails here ; this is shown
by the Artemisise, Cynaraceee, and Astragalacese ; but the
influence of the neighbouring mountains modifies the
character of the vegetation as determined by the climate.
The plants of the steppes are destroyed in the summer
by the drought, whilst Kabarda is well watered from the
Caucasus.

C. Koch remained during October in Ossetia, in the
middle of the high Caucasus, and the offsets connecting
it towards the south with the Armenian highlands, and
then travelled in Imiretia until the end of the year, cer-
tainly too late a period to allow of the botanical character
of the country being completely ascertained. The reports

GIWGRAPHICAL BOTANY. 351

are partly limited to lists of the localities of the autumnal
plants which he was then able to collect. The alpine
flora, even at elevations of 7 SO 00' was but slightly
represented by its characteristic forms (ii, p. 69) ; these
high mountains are altogether more sterile than the Alps,
which the author attributes primarily to the rarity of
glaciers in the Caucasus, as if the alpine meadows of the
Ty r l were only fertilized by melting ice. He then goes
so far as to assert (p. 91), that the disintegrated soil of
Ossetia, the steep rocks and precipitous defiles of this
alpine district, are not adapted to the production of
humus, and that this is the cause of the total absence of a
luxuriant vegetation there. But the author is not clear
upon this point, and does not separate general from local
conditions ; for he speaks at the same time of clay-slate
plateaux, but little supplied with water and destitute of
woods, extending between the defiles and valleys to the
ridge and lateral offsets of the Caucasus. Upon this form
of mountain and peculiarity of soil the alpine poverty of
Ossetia appears to depend ; that it also prevails over the
well- wooded slopes of the northern Caucasus is not pro-
bable. But Ossetia shares this deficiency of alpine
vegetable forms with the mountains of the south of
Europe, where alpine pastures abounding in species are
but rarely developed, and where this phenomenon is occa-
sioned by the deficiency of water upon narrow crests and
summits. Ossetia does not possess the fine forests of
the northern promontories of the Caucasus. Even in
the true forest region there is a perceptible deficiency of
wood, and frequently the soil is scarcely covered with a
scanty underwood : e. g. at Zrchinwall (p. 55) consisting
of Corylus, Cornus mascula, Paliurus, Crattegus, Prunus
insititia, and Jumperus. The traveller only met with
wooded slopes at Dschedschora, in the district of Gudaro,
(p. 82) ; here deciduous trees predominated, and the
Coniferae present were Pinus abies, picea, and orientalis,
Taxus, and Juniperus communis. The deciduous forests
consisted of the oak, beech, rnaple, lime, and alder (Quercus

352 GEOGRAPHICAL BOTANY.

iberica Stev. and Robur (?), Carpinus orientals, Fagus,
Acer platanoides, Tilia parvifolia, Alnus denticulate
C. A. M.) ; the underwood of Euonymus ZatifoZius, Ehamnus
frangula and eatkaftica, Staphylea pinnata, Viburnum
orientate, Argyrolobium lotoides, and Lonicera c&rulea.

The Imiretian slopes of the Caucasus in the upper
valley of Rion (p. 129) are more abundantly wooded;
above the vine mountains of the latter, mixed forests of
deciduous trees ascend to a considerable height ; in ad-
dition to the trees above mentioned, the chesnut, various
fruit trees, and poplars were found at Oni, as also among
the shrubs, Ilex, Azalea pontica, and EJiododendron Cau-
casicum, Rhus Cotinus, together with Smilax excelsa. At
Glola, wild fruit-trees, especially Pyrus communis, and
Prunus avium, extended to beyond 5000'. A region of
subalpine shrubs, of which Arctostaphylos and Azalea
pontica ascend together high up at the glacier of Rion,
immediately succeeds the deciduous forests ; and with
them, subalpine herbaceous perennials, as Aconitum nasu-
tum Fisch., Pyrethrum macropliyllum, Doronicum Cauca-
sicum, <fec. Lower down in the valley of Rion, Koch
describes a fine primitive forest at Kutais (p. 166), con-
sisting of magnificent trunks of Carpinus orientalis, oaks,
and high tops of the chesnut and plane-trees projected
singly ; in thickets, luxuriant lianes of the grape-vine,
Smilax, and ivy, upon the branches of which the mistletoe
grew, and from which Usnete were suspended.

The journey from Tiflis to Erivvan through Georgia
and Russian Armenia was made during the months of
April and May, in 1837, and yielded a rich booty. The
forests of Somchetien differ from those of Imiretia, in
the more regular growth of the trees, and in the absence
of evergreen shrubs and lianes (p. 350). They consist
of Querous iberica and pedunculata, Carpinus Betulus
and orientalis, Acer platanoides sndpseudqplatanus; with
isolated examples of UZmus excelsa Bork., Fagus, and
Acer tartaricum. The soil of these forests consists of a
thick layer of humus, and this black earth produces the

GEOGRAPHICAL BOTANY. 353

beautiful mountain pastures which alternate with the
former. The traveller soon ascended the high mountain
chain between Kur and Araxes, forming the boundary
between Georgia and Armenia (this he calls the lower
Caucasus, Giildenstedt and Klaproth call it the promon-
tory of Ararat), which, according to Parrot's measure-
ment, ascends to an elevation of 12,780'. But vegetation
was still backward in this region, for even in the
Armenian highlands, few of the herbaceous plants and
Grasses which, with a thorny underwood of Tragacanth,
cover the bare heights, were in flower (p. 386). However,
on going from Alagas, near the valley of Araxes, to
Eriwan, Koch was amply compensated by the banks of
the Kasach (p. 397). The climate is so dry, that even
in May the soil is parched and barren, whilst in the more
elevated regions vegetation has scarcely commenced.
But by artificial irrigation, the cultivation of the soil may
be effected even during the hot and dry months, the
fields and orchards surrounding the villages then resemble
oases in a stony desert. Fruit trees were planted generally,
especially peach trees and apricots ; besides these there
was a natural arboreal vegetation along the Kasach valley,
consisting of Elceagnus and Populus, with Prunus incana,
and Tamarix. In Eriwan, especially, the greatest at-
tention is paid to the cultivation of fruits and the vine,
and the traveller had never met with more beautiful
gardens than he saw there.

Schrenck continued his travels in Soongarei in 1843,
and has already made known the plants he found in
that year (Bulletin Petersbourg, iii, pp. 106-10, 209-12,
305-9) . They belong to the following genera : Ranunculus
(2 sp.), Stubendorfia, nov. gen. Crucifer., Isatis, Geranium,
Zygophyllum, Haplopliyllum, Euphorbia (2 sp.), Sophora,
Oxytropis, Astragalus, Seseli, Lomatop odium, nov. gen.
Umbelliferse, Carum, Artemisia (2 sp.), Cliamageron near
Henricea, Saussurea, Cousinea (4 sp.), Plagiobasis, nov.
gen. near the preceding, Jurinea, Serratula, Echinops,
Echinospermum, Eremostachys, Artlirophytum, nov. gen.

354 GEOGRAPHICAL BOTANY.

Chenopod., Pterococcus, Statice, Populus^Ephedra.AlUum,
Typlia, and Triticum (2 sp.)

Middendorf has commenced the arrangement of the
results of his journey through the north of Siberia, which
was mentioned in the preceding annual report (Bulletin
Petersbourg, iii, pp. 150 et seq.) The Tundres of the
Taimyr country, i. e. the peninsula situated between the
lower Jenisei and the Katanga, contain in their diluvial
loam, in addition to the mammalia of the diluvium, large
masses of wood either in a bituminous state, such as is
found in the peat moors, or converted into peat. In such
of these tracts, however, as were beyond the tree limit,
the stems were only met with lying horizontally, and were
compared by Middendorf to the floating timber of the
arctic coast, and from which, by the rising of the land,
they may have gradually attained the interior. The trees
appear to be of the same kinds as those in the forests of
New Siberia and the fluviatile valleys of Siberia, consist-
ing principally of the beech and larch ; they have not
yet, however, been examined microscopically, hence these
statements require confirmation. The climate of the
Taimyr country appeared to be less cold than might have
been expected : from the 6th of June to the 8th of August
there was no frost there; constant fogs and storms
(especially in summer, so that in May, June, and half of
July, the altitude of the sun could only be taken three
times,) indicated great irregularities in the distribution of
heat in the atmosphere. The high surface of the country,
which rises to an elevation of 1000', was perfectly free
from snow in the summer ; even in the winter, storms
sweep the snow into the lowest parts, frequently leaving
the heights bare. In the middle of July, Middendorf
saw at Taimyr 1500 square miles (Eng.) free from snow,
in a few narrow valleys only was any still remaining.
The lakes only freeze to a depth of eight feet ; the layer
of snow then protects them from any deeper penetration
of the frost. As regards the botanical results, we must
wait for further reports, on account of the want of accu-
rate determinations of the plants.

GEOGRAPHICAL BOTANY. 355

The thermometric observations made by Stchoukine,
from 1830-1844 at Irkutsk (1330 English feet above the
level of the sea), give (the months being reckoned as
lunar) the following average of temperature :

January . . - 19'9 c. July . . 18'5 c.

February . . - 13'6 c. August . . + 13'75 c.

March . . - 3'25 c. September . + 6'75 c.

April . . + 575 c. October . . 375 c.

May . . + 12'25 c. November. . 14-25 c.

June . . + 17'6 c. December . . 19'9 c.

Mean temp. = + 0'01 c.

Maximum = + 35 (once in 1843, 39'5 c.)

Minimum = 35

Turczaninow's " Flora of the Baikal Regions" (see the
Annual Report for 1842) has been continued, and has
now reached the end of the Umbelliferae (Bulletin de la
Soc. de Moscou, 1843-44). The following families have
thus far been treated of: 3 Rhamnese, 94 Leguminosas,
69 Rosacese, 5 Onagrariae, 6 Haloragese, 1 of the Cera-
tophylleaB, 1 of the Lythrariae, 2 Tamariscinese, 1 of the
Portulacese, 8 Crassulaceae, 1 Nitraria, 9 Grossularieae, 19
Saxifragese, 48 Umbellifera 3 ,, with the recently separated
genera Physolophium (Angelica saxatilis Turcz.) and Czer-
ncevia (Conioselinum Czern&via F. M.) Altogether 542
Polypetalous plants have now been fully treated of.

Kittlitz's work contains some very interesting illustra-
tions of the characters of the vegetation of Kamtschatka ;
his botanical sketches of the countries, which were made
during the well-known voyage of the younger Mertens
round the world, and described in the text with a perfect
comprehension of the physiognomical characteristics, form
one of the most valuable contributions to botanical geo-
graphy made during the past year (Vierundzwanzig Ve-
getation s-Ansichten von Kiistenlandern und Inseln des
stillen Oceans, aufgenommen in den Jahren 1827-9 durch
F. H. v. Kittlitz. Siegen und Wiesbaden, 1844-5, 4to).
As we cannot omit making a full report upon this work,

356 GEOGRAPHICAL BOTANY.

we shall preserve as far as possible the excellent language
of the text which accompanies the copper-plates; they
afford a sample of the author's power of observation.

The physiognomy of Central and Northern Europe
agrees with that of Kamtschatka much more completely
than we should anticipate, considering the great difference
between their longitudes : the number of European plants
is very considerable (p. 53). The peninsula is divided
into an eastern and western half by its mountain-chain,
In the former, rise the conical volcanic mountains, of
which the Kliutschewsk, according to Erman, is 14,800'
in height, or as Kittlitz expresses himself, they rival the
Peak of Teneriffe in height, and excel all other volcanoes in
the perfection of their conical form. They alternate with
long mountain -chains whose rugged tops are covered with
snow, whilst the remainder of the district is adorned with
the growth of noble forests and pasture. On the west side,
however, the coast is low and marshy, passing towards
the interior into a broad plain of fertile land, the soil of
which is watered by numerous streams, and is covered
partly with woods, partly with luxuriant grassy plains in
their original and natural state, For the purpose of car-
rying out this sketch completely in detail, the author has
given five tables, which indicate the botanical character
of the forests and grassy plains in the summer months
(July to September).

Grass Plain at Awatscha, therefore in the neighbour-
hood of Peter-Paul's harbour (plate XVII). This picture
represents a luxuriant woody prairie, abounding in plants,
and containing scattered groups of shrubs, and the open
surface of which is inclosed by a wood of birch (Betula
Ermani). This birch is the principal forest tree of the
country; it somewhat resembles the oak in the knotty
and flexuous growth of its stem, and differs moreover
from Betula alba in its bark, which is gray and much
torn, whilst the leaves agree with those of the common
birch. A thicket of alders and willows denotes the vici-
nity of the stream ; some of these are shrubby, others

GEOGRAPHICAL BOTANY. 357

tall in growth, resembling that of the poplar, and with
these woody plants the gregarious Spiraea Kamtschatika
(Schalameynik), a plant which throughout the summer
characterises Kamtschatka above all other countries, and
here repeats the Panax-form of the north-west of America
in a physiognomical point of view : " A plant of wonder-
fully rapid growth, which in a few weeks acquires a height
of more than 10 feet, whilst in the autumn it disappears
still more quickly, without leaving a trace, for a single
frosty night is sufficient to cut it off to the ground."
Above the large, crenate leaves, the stems display in
July their white bunches of flowers, which subsequently
acquire a gray tint. Single plants of a very tall Hera-
cleum (H. Panaces ?) grow among the Spiraeas, from the
juice of which the natives prepare sugar. The grass
covering these prairies attains an astonishing height ; at
first, indeed, it is overshadowed by shrubs of Cratcegus
and Salisc, with thick stems, which project here and there,
but these at a later period scarcely extend above the
rapidly developed culms of the grass. The same applies
to the herbaceous perennials, which are mixed in large
numbers with the Grasses, and of which the following
are mentioned : 2 Sanguisorbtf, Angelica, Epilobium an-
gmtifolium, Senecio cannabifolius, Cacalia hastata, 2 lilies
with large orange flowers (one with stems of the height
of a man, probably L. Kamtschatkense Lour.), and Fri-
tillaria Kamtsc/iatkensis, the latter under the name of
Sarannah. Of these, Senccio and Epilobium are the prin-
cipal ones which contribute to the physiognomy of the
land. The former, although as high as a man, is laden
with flowers, and frequently colours the surface of the
meadows of a pure yellow colour, whilst the latter pro-
duces a splendid red. The Sarannah, which is every-
where met with in short grass, yields in its tubers an
excellent article of food, which, although difficult to dig
up, often supplies the place of bread.

Plate XVIII leads us to the Forest on the Upper
Kamtschatka river, which, lying in a valley running Ion-

358 GEOGRAPHICAL BOTANY.

gitudinally towards the east, traverses plains that are
extensive towards the north, and almost everywhere
wooded. Here, but here only, a different kind of birch
constitutes the predominant forest tree, which the author
regards as one of the European species, and denominates
Betula alba (B. pubescens of Erman). It is so distinctly
separated geographically in the neighbourhood of the
river from B. Ermani, that on the road from Ganal to
Puschtschina, whilst from the coast to this place the latter
only is met with, the white birch suddenly begins to
form the forests as soon as the upper course of the Kam-
tschatka river is reached. Together with the birch, we
here find drawn a group of tall balsam poplars as straight
as a line ; this tree by itself forms large woods in the
middle of Kamtschatka. The underwood and shrubs
consist principally of Spircea, next of Lonicera, Crat&yus,
Prunus, and Salix. In the glades, in the midst of scanty
grass, a dark blue Iris grows ; it is everywhere common,
forming an incomparable ornament to the country, and is
succeeded at a later period by several Synantheraceous
perennials with beautiful flowers, as Aster, Achittea, and
Sonchus Sibiricus.

Forests of Central Kamtschatka (pi. XIX, XX). A
strip of land extends across the middle of the peninsula,
from the west towards Cape Kronotzkoi; it is wooded
with Coniferse, no trace of which exists in the other dis-
tricts. The forests consist of two kinds of fir-trees, the
larger of which resembles the Canadian larch, the other
has the growth of our red pine, with which it is probably
identical : here also the birch and the aspen are associated
with them. As the pine-forests of Kamtschatka differ
from those of the north-west of America in their dryness,
so also the underwood merely consists of a thicket, 3 feet
high, composed of Roses and LonicercB, and beneath them
again a large number of bacciferous plants are concealed,
Vaccinia, Rubi, and Empetrum, exactly as occurs under
similar conditions in Scandinavia, so much so, that even
the species mentioned of these genera are identical with

GEOGRAPHICAL BOTANY. 359

them. Among the edible fruits, Rubus arcticus has the
most agreeable taste ; the elongated dark blue berries of
a Lonicera come next, their taste is not inferior to that
of the finest cherries, and they are prepared with milk or
Sarannah to form a favorite article of food with the natives.
The Kamtschatka river is constantly changing the course
of its valley, and hence, like the rivers of Russia, its banks
are steep (Jar) on the side excavated by the current, whilst
sandbanks (Pessok) are deposited by the water on the
opposite side. On the former, the old pine-forest extends
down to the river, and by the falling in of the banks is
carried away as floating timber ; on the latter, different
woody plants have settled, the period of formation of
which is later than that of the former : first, thickets of
willows, then larger deciduous trees, willows, alders, and
poplars appear to follow. The difference in the foliage
commonly expresses a difference not of the age of the trees,
but of the period at which the district became wooded.

Mountain-Forests of the Eastern Coast (pi. XXI), ex-
tending over its steep declivities. These forests, which
are also composed of Betula Ermani, and sometimes con-
tain tall trees of Sali%, appear far lighter than those in
the fluviatile valleys ; but the thickets of underwood and
shrubs extending between the trees are proportionately
thicker, and contain a larger number of plants. This
character is evident, even at a level of 500',, and ex-
tends high up the mountain. But at a greater elevation,
the birch trees gradually diminish in number, preserving
the same state of growth, until at last they disappear, and
give place entirely to the shrubs, just as the latter are
displaced by the alpine flora, according to the same law.
These thickets of shrubs are in general impenetrable to
man, and represent the pine-region in Kamtschatka.
They consist of Pyrus sambucifolia Cham., Alnus incana,
and a pine which is probably a variety of Pinus Cembra,
and is called Kedrownik. The former of these shrubs
predominates in the lower regions, and disappears at an
elevation of 1000'. The Kedrownik grows even in the

360 GEOGRAPHICAL BOTANY.

vicinity of the coast, but appears to be most widely dif-
fused between 1000' and 2000'. Its nuts are nutritious,
and are eaten, as is also the fruit of Pyrus mmbucifolia:
The most extensive thickets consist of the northern alder,
which also grows in the lower regions, in common with
two others, but between 2000' and 3000' exists alone,
limited by the alpine flora, bare stones, and eternal snow.
On all the high mountains of Kamtschatka there exists a
region in which it exclusively covers the soil. Its upper
limit had been previously determined correctly by Erman
to be 2890', i. e. more than 2000' beneath the snow-line
(5000') ; but Salise arctica (4974'), Parry a Ermani, and
Saxifragi MerJcii ascend as high as the latter.

Grass Plain in the West of Kamtschatka, on the
Bolschaja-Reka (pi. XXII). The south-western slope of
the peninsula is comparatively poor in pictorial beauty
and botanical variety; the forest-growth is less than in
the east, the morasses are more extensive, and bushes of
willow predominate almost everywhere with the peat-
moors. The landscape, which was taken in September,
is remarkable from the astonishing height of two wither-
ing Umbelliferous plants, which give a most peculiar
character to the grass plains of the west. They probably
belong to the genera Angelica and Heracleum ; their
strong stems appear more than fifteen feet high ; thus,
growing in numbers, they project far beyond the Grasses
and other herbaceous plants. We next have a tall gre-
garious Urtica, 10' high, and from which the natives
prepare a valuable yarn. The remaining plants agree
generally with those of the grass plains in the east.

The Algae of Kamtschatka are described and figured in
the splendid illustrated work of Postels and Ruprecht ;
they were also collected in the expedition of the younger
Mertens (Illustrationes algarum in itinere nauarchi Liitke
collectarum. Petropoli, 1840, fol.)

Zuccarini has published a very valuable sketch of the
Flora of Japan (Notizen liber die Flora von Japan und
die bisher hieruber vorliegenden wissenschaftlichen Lei-

GEOGRAPHICAL BOTANY. 361

stungen : in the Miinchener gelehrte Arizeigen fur 1841
and 1844, id. pp. 430 et seq.)

It must first be remarked, in regard to the notice
given in the Annual Report for 1843, of the progress of
the author's Flora Japonica, that this work has indeed
experienced an interruption, but that by the completion
of the part which treats of the Conifers, the number of
this order found in Japan has been increased, far beyond
that previously given, i. e. to 30, which are distributed
through 14 genera. Zuccarini's present work contains a
catalogue of all the genera as yet known in Japan, with
the number of species in each family. The latter amount
in all to about 1650 species; but as Zuccarini estimates
the number of the Japanese plants contained in the
herbaria of the Netherlands at 2400 species, the sta-
tistics must ultimately be altered in proportion as the still
remaining families in V. Siebold's work are worked out.
With the proportional numbers of the genera and families
this will not be so much the case; hence Zuccarini's sketch
acquires a permanent value. He enumerates the following
as the most remarkable general results of his investiga-
tion : 1 . The large number of families of plants repre-
sented in Japan, of which, according to Endlicher's system,
there are 172. 2. The large number of genera in pro-
portion to the species, for 621 are already mentioned in
the catalogue, and probably 700 are contained in the
herbaria (it must, however, be remarked, that Zuccarini
has included the Chinese genera found in Beechey's
voyage, as also those from the Bonin Islands). 3. The
limitation of endemic genera to a single species, corre-
sponding to the monotypes of the Canary Isles; a condition
which applies to the greater part of the new genera from
Japan, whilst the remainder contain at present only two,
or, at the most, four or five species, and some mono-
types also of North America and India, and the European
Humulus, possess in Japan a second, but only a second
species. 4. The very large number of woody plants in
so high a latitude, both from woody families belonging

362 GEOGRAPHICAL BOTANY.

to the temperate as well as the tropic zone, from the latter
of which representatives of the Palms, Pandaneae, Lau-
raceae, Ternstroemiaceae, &c., together with numerous
bamboos, are here in part diffused further towards the
north than in other meridians of the northern hemisphere.
5. The endemic character of the flora of Japan, which is
not connected, like Siberia, with that of Europe, having
but very few species in common with Europe. We have
not space, unfortunately, to enter more minutely upon
the consideration of the components of the catalogue of
the genera ; we shall, therefore, merely confine ourselves
to the mention of those families which are remarkable
from the number of species they contain, and to the enu-
meration of some of the characteristic botanical forms of
Japan. The predominating families are : Synantheraceae
(124), Graminaceae (90), Rosaceae (90), Leguminosae
(72), Liliaceae, in the extended sense (60, of these 25 are
Smilaceae), Cyperaceae (48), Labiatae (47), Ranunculaceae
(42), Umbelliferae (40), Amentaceae (38), Orchidacege (35,
principally of North American and European genera),
Ericaceae (36, of North American genera), Coniferae (30),
Urticaceae in the extended sense (about 30), Cruciferae (30).
Characteristic forms (excluding several tropical representa-
tives): Melastomas(4), Zanthoxyleae (6), Aurantiaceae (10),
Ternstrcemiaceae (19), an Opuntia, the source of which is,
however, doubtful, Magnoliaceae (10), one of the Proteaceae
(Helicia Z .) , Lauraceae (18,) Palmeae (4) , Musaceae (4 Musa] ,
Scitamineae (7), the Haemadorous Aletris, Dioscoreae (5), 1
PhilJiydrum, Commelyneae (5), Eriocaulon (4), and Cycas (1) .
Zuccarini and v. Siebold have described some new
genera of Japanese plants in the memoirs of the Academy
of Bavaria (Plantamm quas in Japonia collegit de Siebold
genera nova. Ease. 1, 1. c. iii, pp. 719-49). List of these
genera : Pityrosperma (a Ranunculaceous plant with 3
species, one of which is Actcea Japonica Thunb.) ;
Pteridophyllum (connecting link between Hypecoum and
Fumaria) ; Eucapnos (Diclytra spedabilis D. C.) ; Tro-
chostigma> with 5 sp. (probably the type of a new family

GEOGRAPHICAL BOTANY. 363

allied to the Ternstrcemiacese) ; Corchoropsis (one of the
Tiliaceae) ; Tripetaleia (doubtfully placed among the
Oleaceae) ; Stephanandra (affinity also doubtful, probably
belonging to the Rosacese) ; Ceraseidos (one of the
apetalous Amygdalaceae plants) ; Platycaria (one of the
Juglandese) ; ScMzocodon (Polemoniaceae) ; Conandron
(allied to Ramondia) ; PhyllostacUys (Bambuseae). Accord-
ing to the author, the bamboo-stems of commerce, as also
the pepper-canes as they are called, are obtained from the
Bambusese, which are common in Japan, and of which
there are 1 5 ; they seldom, however, flower, and therefore
the species are but imperfectly known.

Royle has drawn up some remarks upon the vegetation
of Afghanistan, Cashmere, and Thibet, from the truly
very inconsiderable collections of Vigne (Travels in
Cashmere, Ladak, Iskardo, &c., by G. T. Vigne, 2d edit.
London, 1844, 8vo, Appendix). However, these frag-
mentary reports are of interest, on account of Royle's
intimate acquaintance with the botanical character of the
Himalayan mountains, the use he has made of other
sources, and the general plan of his investigation. Thus
he starts with the question of what constitutes the northern
and western boundaries of the indigenous plants of the
Indian plains. He considers it as an established fact,
that the western boundary of the Indian flora along
the Indus is formed by the Soliman mountains, and, in
fact, the influence of the monsoon and summer-rains,
upon which the vegetation of the tropical plains is de-
pendent, disappears entirely in the district of this meri-
dional chain, on the line from Kelat to Peshawar. Rovle

is especially indebted for the observations upon the western
localities of Indian plants to the traveller Falconer, who
is now his successor in the Botanic Garden at Saharun-
pore. The latter found Butea frondosa even on the
Jhelum, the most westerly of the Punjab rivers ; the
Chenopodese of northern India accompanied it as far as
Peshawar. Above Attock, on the Indus, the charac-
teristic plants of the British Himalaya again recurred.

364 GEOGRAPHICAL BOTANY.

Even from Attock, according to Elphinstone (Cabul, p.
130), the tropical rains extend northwards as far as
Hindu-Rusch, without the high flats of Afghanistan being
moistened by them; for Surat would there form its
western boundary, at which place, in summer, whilst e. g.
rain still falls in Pukkely, the sky is overcast for a month
only, and merely occasional showers fall. Thus the
double harvests of the Indian year, which are occasioned
by the rainy season, cannot be obtained west of Jellalabad
(Irvine, Journal of As. Soc. of Bengal). Hence between
Jellalabad and Gundamuc, on the road to Cabul, the
periods of development of the vegetation are suddenly
changed. " In Gundamuc," writes Burnes, " the willows
flowered at the end of February. On the llth of March
the first spring flower appeared ; it was a sweet-smelling
Iris. The apricots began to unfold their buds on the 1st
of April; the wheat here was three inches above the
ground, whilst in Jellalabad it was already cut." But when
we take into consideration the elevation of the soil above
the Indus and its tributary streams, it appears clear that
the tropical conditions of the vegetation only extend so
far west in the valleys. In fact, Royle does not allude to
the important question, to what elevation the mountain-
slopes which limit these fluviatile valleys on every side are
reached by tropical rains ; but as regards Cashmere, a
valley lying far to the east of Peshawar, we know that the
atmospheric precipitations of the spring cease to occur at
that period at which the rainy season commences in the
Indian plains and the low valleys of the Himalaya. Thus
it appears, from all the descriptions, that the more elevated
regions in the neighbourhood of Attock and Peshawar
are not subjected to the monsoon. This explains Elphin-
stone's statement, that a number of English plants thrive
in the gardens at Cohaut, where plum arid peach trees
were in flower at the end of February, and where weep-
ing willows, plane, and apple trees were thriving upon
European meadow plains. From these reports, it is pro-
bable that the entire district west and north of the

GEOGRAPHICAL BOTANY. 365

Jhelum, or of the salt-chain, which is intersected by in-
numerable offsets of the Himalaya and Soliman moun-
tains, with the exception of the lower fluviatile valleys, is
free from all those Indian vegetable forms which, up to
the foot of these mountains, are extended in an uninter-
rupted distribution over the Punjab.

But Royle's investigation passes over unnoticed a still
more important aspect of the question regarding the
boundaries of the Indian flora. Hitherto we have only
treated of tropical forms of vegetation, to the growth of
which the rainy season is unfavorable ,- but in addition to
these, India possesses in the Himalaya and the monsoon
region that mountain vegetation also, in which the European
type is repeated, Here the question arises whether the
areal limits of the latter are the same as those of the
former, with which, in fact, they partly grow in common
on the western chain of the British Himalaya, without,
however, being favoured in the same degree, during their
period of vegetation, by the tropical rain. The knowledge
of this remarkable coexistence of the productions of two
climates, for which we are also principally indebted to
Royle's former investigations, has not induced him to
devote his attention to the question of whether there are
not forests of Himalayan trees in other regions which do
not shade tropical plants in the rainy season. However,
the simultaneous publication of Jacquemont's Journal at
Cashmere has thrown some light upon this obscure point
(Voyage dans 1'Inde, vol. iii, p. 169). The traveller
describes his journey from the Punjab to Cashmere over
the Pirpanjol, the Himalayan Pass, which Royle, relying
upon Bernier's descriptions, had formerly marked as a
sharply-defined limit of the vegetation of the Indian
flora, which assertion he now himself withdraws pretty
openly. During the ascent, the pomegranate and olive
trees disappeared at an elevation of 16 1700m., and
soon after, Pinus longifolia also. A region of oaks,
Pinus attenuata, and firs was next met with, which, on
the northern slope of the chain, extended above the level of

p66 GEOGRAPHICAL BOTANY.

the Pass (2681m.), whilst on this side it terminated below
the alpine meadows. The alpine vegetation presented
merely local differences from that of the British Himalaya ;
its spring-plants were in flower at the commencement of
May. On the north side, therefore, Jacquemont first
met with the same trees he had left on the southern
slope, and further down, in the district of the valley of
Cashmere, he arrived at forests of an ^Esculus of the
same species as that indigenous to the British Himalaya.
The older opinions concerning the Pirpanjol, which Royle
disseminated, are contradicted by these Reports. But as
there is no tropical region in the valley of Cashmere, we
have here also a proof that the diffusion of the Himalayan
plants is not limited by the boundaries of the monsoon.
The tropical forms of India may be wanting in Cashmere
and there is no evidence to show that they exist there
and yet the forest trees may appear the very same, and
the character of the vegetation for the most part identical
with that of the British Himalaya ; in fact, the greater
number of species may be common to both of them. The
natives of the Pirpanjol say that it is always raining there
(p. 225) ; hence this Pass may form one of the points of
the boundary, as far as which tropical forms accompany the
wooded slopes of the Himalaya. When thus considered,
all the known facts are connected under a common point
of view, but they are by no means sufficient for deter-
mining the absolute sphere of diffusion of all the Indian
plants. Although Royle has rendered it probable that this
area does not extend west of the monsoon-limit, yet the
line at which the Himalayan plants cease, towards the north,
is either totally unknown to us, as is the case beyond
Cashmere, or merely indicated by uncertain evidence.

Royle's statements regarding the flora of the elevated
plains of Afghanistan are very general ; but where Griffith
is his authority, the fragmentary notices derived from his
letters are substantiated by the catalogue of a series of
Afghanistan genera, the seeds of which were also trans-
mitted by Griffith. They are nearly all European forms,

GEOGRAPHICAL BOTANY. 36'$

and principally the following : Aconitum and Papaver ;
5 European Cruciferse and Tauscheria ; Silene and Are-
naria ; Euta and Peganum ; Euphorbia and Phyllantlius ;
several Astragalaceae and Caragana ; Rosa and Crat&gm ;
Epilobium ; Prangos pabzdaria ; several Carduacece with
Centaur ea and Cicorium ; Campanula; Heliotropium and
Onosma ; Pedicularis, Linaria, Veronica, and Verbascum ;
Hyoscyamus, Samolus, Plantago, Hippophae, Humex, and
Polygonum ; Blitum ; Iris and Tulip a. Irwine treats fully
of the cultivated plants of Cabul (loc. cit.) Wheat, barley,
lentils, and peas are sown ; they are protected during the
winter by a layer of snow, and harvested in June. To the
summer crops, which usually require irrigation, belong
Phaseolus radiatus, Cicer arietinum, Panicum miliaceum
and Italicum, maize, and rice ; these are sown in May, and
harvested in the months of August and September.
Besides the European vegetables, Solanum melongena and
several Cucurbitacese are cultivated, which require much
manure and water. The meadows yield abundant crops
of hay, and contain some excellent species of trefoil : one
of these is denominated Trifolium giganteum ; Medicago
sativa is also widely diffused. The fruit trees of Cabul are
celebrated : in addition to the fruits of central and eastern
Europe, those of El&agnus (Sinjet and Sinjilla) and
jEdgeworthia buxifolia, one of the Theophrasteae, are
mentioned.

Falconer discovered, in Cashmere, the plant yielding
the Costus of the ancients, a substance which still forms
an article of commerce in India, under the name of Koost
or Koot. It is obtained from the aromatic root of a new
alpine species of Carlina (AucMandia), which Falconer
has accurately described (Linnsean Trans., xix, p. 23).
He has also proposed there (p. 101) the Asclepiadaceous
genus Campelepes, from Peshawar. Falconer's so-called
Fothergilla, which forms large bushes in Cashmere, and
the wood of which, according to Yigne, is called Chob-i-
pan, is a new type of the Persian Parrotia (P. Jacque-
montiana Decs.)

; GEOGRAPHICAL BOTANY.

From the elevated valley of Astore, between Cashmere
and Thibet, Vigne brought the following plants : Aconi-
tum heteropliyllum, Anemone discolor, Podopliyllum, Dian-
thus, Geranium, Epilobium, several Gentians, Swertia and
Ophelia Chirata, Polemonium cceruleum, and Dracocepha-
lum Royleanum. Here, far above the tree-limit, we find
the elevated plain Deosuh, at an altitude of 13,000', the
soil of which is rendered verdant by dwarf- willows and
alpine herbs, whilst the valley in which the Indus runs
in Thibet is bare, a few plants occurring only at the
snow-line. Falconer found here a new Rheum and two
species of Pyrola, which, as Royle remarks, are the only
Ericaceous plants in Thibet. Vigne' s plants from
Iskardo agree pretty accurately with the older collections
from Kunawar : Actaa, some Cruciferae, Silene Moor-
croftiana, Acer microphyllum, Myricaria, Biebersteinia
odora, Astragalacece, several Potentillce, Saxifraga steno-
phylla, Hippophde and Salsola.

Jacquemont's work on his travels, which has been
mentioned above, is now complete, and affords extensive
contributions to our knowledge of India in a botanico-
geographical point of view, especially the flora of the
British Himalayas and those of Thibet (Journal, vol. i-iii.
Paris, 1841. Vol. v, Descriptions des Collections. Ib.
1844, 4to. 2 vols. plates). The admirably-kept journal
of this traveller, which is printed unaltered, contains, of
course in a fragmentary form only, the impressions pro-
duced by the character of the vegetation of the Himalayas,
and separate regions of India ; but in the last section of
the work, the more rare and new plants of Jacquemont's
herbarium are treated in systematic detail by Cambes-
sedes and Decaisne, and illustrated with 180 plates.

In Lesser Thibet, J. travelled on the road to Ladak, in
the valley of Spiti, as far as Danker, where at an ele-
vation of 17,000', at the limit to vegetable life, he found
the new Anthemideous genus Attar dia, a Nepeta, and
an Urtica. The villages in the valley of Spiti, according
to Jacquemont, are situated on a higher level than that

GEOGRAPHICAL BOTANY. 369

formerly stated by Royle, e. g. Nako at 3658m. ; and
the cultivation of the Cerealia, which is limited to Hor-
deum hexastichon and celeste, and a Panicum, extends
here to 3962 m., whilst in the southern Himalaya it only
extends to 3048 m. Woody plants are not entirely
absent from this elevated valley ; even low trees, an
indigenous Juniperus, and cultivated poplars and willows,
are met with. The character of the vegetation, however,
lies in the bushes, which was also noticed by Moorcroft
These consist not only of thorny Astragali, but also of
Genista, Rosa, Ephedra, and Juniperus. The absolute
limit of elevation of the Phanerogamia west of Bekar was
most accurately determined by Jacquemont. Here, in
two passes leading from Thibet, Gantong (5486 m.) and
Kimbrong (5581 m. according to Gerard's measurement),
he left these plants below him. The leguminous shrubs
of the valleys of Kunawar and Lesser Thibet were not
found on the slopes of this pass, only a few alpine plants,
the last of which was met with at Gantong, about
2 300 m. beneath the summit, hence at a level of
5200 m. Here he found two Potentill<B, Corydalis pliy-
socarpa, the new Caryophyllaceous plant Periandra COBS-
pitosa, which resembles in appearance Silene Acaulis,
with Allardia and JEntrichium Jacquemontii (Decs, iij
p. 309). Much lower down, the traveller met with a
rose, forming the last shrub, and considerably lower still
a Juniperus. At Kinbrong the vegetation also disap-
peared 300 m. below the pass, with a Ranunculus, Cory-
dalis, and Ligularia nana ; but at a level of 5400 m.
Jacquemont saw an isolated green spot in the stony desert-
waste. This was the highest evidence of vegetable life
which he perceived (ii, p. 298). He estimated the snow-
limit here at little less than 6000 m., so that between
the last plants and perpetual snow there is an intermediate
bare region extending through about 2000'.

As regards Kunawar, that remarkable transition-dis-
trict between the British Himalaya and Thibet, on the
central Sutlej, where the influence of the monsoon on

370 GEOGRAPHICAL BOTANY.

the seasons ceases, and the dryness of Thibet commences,
Jacquemont's botanical observations agree with the more
copious reports of Royle. The forests are very incon-
siderable, the growth of grass poor, and kept down by
Tragacanth-shrubs {Astragali} , which are distributed as
far as here; the alpine flora is also very scanty
(ii, p. 269). Jacquemont devotes particular attention to
the cultivation of the grape-vine, which is confined to
this part of the Himalaya, not extending beyond the limits
of the tropical rain (ii, pp. 416 et seq.) Although the
grape-vine is cultivated at an elevation of 10,000', this
is only the case in the bottom of the valley, not on the
mountain-slopes, for it only there receives the reflected
rays of the sun, which are necessary to ripen the grapes,
and there it is also protected from that radiation of heat
which exerts too powerful an effect in cooling the earth
on mountains. Moreover, even in the valley of the Sutlej,
irrigation is indispensable to this branch of culture ;
but although the grapes under these circumstances mostly
ripen well, they are usually dried in the sun, and used
to make raisins, as the wine does not keep long, and
even when new was found almost undrinkable by the
Frenchman. We find the grape-vine as far upwards as
Nako, in the valley of Spiti, and downwards as far as the
mouth of the Buspa, where the climatic line above men-
tioned lies, and where the Sutlej intersects the high
southern chains of the Himalayas.

The chains of the Southern Himalayas, which are
situated immediately opposite to the plain of the north
of India, do not possess any of that variation of soil, by
means of which their vegetation might equal the flora of
the Alps in variety, notwithstanding the mixture of forms
of Tropical and European plants. Plane surfaces are
scarcely anywhere found; and as we have already re-
marked, the broad valleys of Cashmere and Nepaul form
exceptions to the mountain-character. Perpendicular
precipices are also absent. We find everywhere vast in-
clined plains, and the mountain-stream usually entirely

GEOGRAPHICAL BOTANY. 371

fills up the bottom of the valley. Jacquemont says
(ii, p. 130), "the vegetation which covers the inclined
soil is as uniform as this conformation. Variety of
localities causes a region to abound in plants, but here
all the localities are alike." In the upper regions the
forests are generally thin, and belong principally to the
valleys. On viewing from a distance one of these im-
mense declivities, on which there are scarcely any forests,
we perceive lines of a darker green accompanying the few
rivulets which water the mountain-slopes, at great dis-
tances apart. Between them the green is uniformly
pale, for neither meadows nor mountain-pastures thrive
there ; but, with the exception of the summits of the rocks,
an irregular and unfruitful growth of plants prevails
among the blocks and the crumbled portions of rocks.
High mountains occur, which, from the valleys to the
crests, are covered with this mixture of rocks and plants
only. More commonly a thin forest is distributed over
a soil of this kind, between 6000' and 7500', consisting
either of pine trees on the southern declivities, or the
oak, with Rhododendron arboreum, on those which are
colder. It is only at the foot of the mountains that dense
forests, such as those on the Alps, flourish. The elevated
forests of the Coniferous region of the Alps are not met
with on the Himalayas.

At Massuri, Jacquemont measured the lower limit of
the oak forest containing Rhododendron arboreum, and esti-
mated it at 1768 m. (ii, p. 52). This measurement is
tolerably near that given by Royle, who, in this district,
determined the elevation of 5 000' to be the level at which
the forms of the European forests appear in the place of
tropical trees. In his ascent of the Kedarkanta, in the
district of the source of the Jumna, Jacquemont also
estimated the upper tree-limit at 3500 m. (ii, p. 127).
The pine forests (species of Abies) terminated here, and
were succeeded by a shrubby formation of Rhododendron
(probably Eh. lepidotum Wall.) ; where this also disappears,
the alpine soil is covered with turf, consisting of Grasses

372 GEOGRAPHICAL BOTANY.

and Carex, among which Ranunculacese most commonly
spring up, with Iris, Corydalis, and Phalangium. The
above measurement of the tree-limit appears to deserve
the more confidence and to form an indication of climatic
conditions, inasmuch as on the Kedarkanta the soil and
inclination of the summit were favorable to forest growth.
Towards the end of his extended tours through the
East Indian peninsula, Jacquemont's attention was drawn
to an important peculiarity in the progress of the vegeta-
tion on the eastern coast of the district of the Ganges
(Hi, p. 550). In Bengal the soil remains green through-
out the year, because the water flows off these plains so
slowly, that it is retained deep in the soil during the dry
season ; also because in the winter dense fogs, and in the
hot and dry months of spring, transient thunder-showers
occur. Thus, when the traveller landed, on the 5th of
May, at Calcutta (therefore on the coast), the turf was
just as green as at the period of the heaviest precipitations
in August. The treeless country of Puna, in the western
Ghauts, however, in 1832 remained perfectly arid and
parched, even in the latter third of June, just like the
soil of the steppes ; the surface of earth was without a
trace of moisture, and, as it were, glowing in the sun's
rays. Yet on the 1st of July the whole country was green,
even the barest rocks had become covered with turf with
wonderful rapidity. Hence the character of the mon-
soon flora is much more distinctly stamped here than
at Calcutta : but the Bengal coast is anomalous in this
respect. In the greater part of India, the vegetation of
most of the plants is interrupted for a longer period by
the dry season, than in Europe by the winter. The large
shrubs, the sugar-cane plantations, and the turf of Panicese
wither and dry up in November, and their vegetative life
is not again aroused until June or July of the following
year. At Puna the rainy season then lasted but little
more than three months, and ceased at the beginning of
November; but that year threatened to be unproductive,
in consequence ' of too small an amount of rain having
fallen.

GEOGRAPHICAL BOTANY. 373

In the descriptive part of Jacquemont's work, which,
arranged in accordance with De Candolle's system, is
worked out by Cambessedes as far as the conclusion of
the Rosacese, and the remainder by Decaisne, in addition
to a large number of new species, the following genera,
mostly from the Himalayas are proposed: Christolea and
Donepea (Crucifera?), Oliyomeris (Resedaceae), Periandra
(vid, sup.), Anquetilla (Xanthoxylaceae) Leptopus (near
Phyttanthus)> Allardia (v. s.), Melanoseris (Cichoracese),
Belenia (Solanege), Dargeria (Scrophulariaceae), Lasiosi-
phon (GnidicB sp. plures) Girardinia (Urtica sp.), and
J)iplosiphon (a remarkable Epigynous and Monocotyledo-
nous water-plant, the natural affinity of which is not
determined).

The continuation of Bentham's work upon the Indian
and African Leguminosoe, which was noticed in last year's
Report, includes about a hundred Genistas, most of them
from the Cape (London Journal of Bot. iii, p. 338-65).

The new parts of Kortlial's Monographs on the Flora
of the Indian Archipelago (Annual Report for 1841), con-
tain the Melastomacese, Oaks, and the following genera :
Cratoxylon and Tridesmis, Hippocratea and Salacia, and
Maranthes ; Boschia, nov. gen. (Sterculiaceae),, Ompho-
carpus, n. g. (near Grewia), Paravinia, n. g., and Clei-
socratera, n. g. (Rubiaceae). De Vriese has described a
Casuarina (C. Sumatrana J.) found by Junghuhn in
Sumatra in v. d. Hoeven's Tijdschrift (1844, p. 113), also
some Javanese plants (id. p. 336-47) ; the only new plant
is an jffischynanthm. New contributions by Hasskarl on
various families of the Javanese flora are published partly
in the same Journal (p. 49, iii ; pp. 178-228), and partly
in the Ratisbon Flora (1844, pp. 583 et seq.) Montagne
has described some new Javanese Mosses (London Journal
of Botany, 1844, pp. 632-4). Dozy and Molkenboer have
commenced an illustrated work on the Mosses of the Indian
Archipelago (Musci frondosi inediti Archipelagi Indici,
Ease. I. Lugdun. Batav. 1 844). The preliminary diagnostic
characters of about 75 new species have been published by

374 GEOGRAPHICAL BOTANY.

them in the 'Annales des Sciences Naturelles' (1844, ii,
p. 297-316); among these are the new genera Crypto-
carpon, Endotrichon, and Sympliysodon.

An extremely important systematic and illustrated
work, on the Flora of Java, which is now concluded, is
that published by Bennett and R. Brown from Horsfield's
herbarium (Plantae Javanicse rariores descriptse inconibus-
que illustrate. Descriptions et characteres plurimarum
elaboravit I. Bennett, observationes structuram, et affini-
tates praesertim respicientes passim adjicit, Rob. Brown,
pt. i, Londini, 1838 ; pt. ii, 1840; pt. iii, 1844). This
work contains 45 plates, and the following new genera:
Sclerachne and Polytoca (Graminacese), Hexameria (Or-
chidacese), Cyrtoceras (Asclepiadaceae), Stylodiscus (An-
draclme trifoliata Roxb.), Euchresta (Andira Horsjieldii
Lesch.), Mecopus and Phylacium (Leguminosse), Saocope-
talum (Anonacese), Lasiolepis (near Harrisonia Br.),
Pterocymbium (Sterculiacese), and several types from
other countries, which are elucidated in these copious
disquisitions.

Junghuhn's diaries of his travels in Java, which have
been already alluded to, were indeed first published, with
additions, in 1845 (Topographische und naturwissenschaft-
liche Reisen durch Java, von F. Junghuhn, herausgege-
ben von Nees v. Essenbeck. Magdeburg, 8vo), but for the
sake of connecting them with the preceding Annual Report,
we think it better to report upon them now. In the western
portion of the island, as at Gede, the traveller found the
mountain-ridges covered far and wide on both slopes with
Rosamala-forests, i. e. with Liquid-ambar Altingiana Bl.,
the stems of which are recognised even at a distance by
their tall straight growth and white colour, and which
overshadow a thicket of Scitaminese, Melastoma, Rubus,
and other shrubs (p. 165). A rich red soil here covers
almost the whole of the trachyte of Gede. According to
several measurements, the region of the Rosamala-forests
is situated at a level between 2000' and 4000' (p. 436) :
this tree, which is confined to the west of Java, occurs

GEOGRAPHICAL BOTANY. 375

singly as high as 4500', and as low as 1500'. It is one
of the most gigantic formations of the vegetable world,
and attains on an average a height of 150': the stems
when cut down measure 15' in circumference, 12' above
the root; their length below the point at which they
branch amounts to 90' 100', and the crowns extend to
a height of from 50' to 80' beyond this. Cocoa palms
would scarcely reach as high as these crowns. Above
the Rosarnala-forest on Pang-Gerango, came forests of
Laurineae, Castanea, Oaks, Schima, and Fagraa, which
were far more abundantly filled with climbing plants
(e. g. Freycinetias and Calamus] and parasites (Orchidacese
and Ferns) ; and these again were succeeded by the Podo-
carpeae. But even beyond the limits of Podocarpus the
arboreal form is not wanting here, as is the case on other
mountains. On the summit of Pang-Gerango itself, at a
level of 9200', Tkibaudia vulgaris^. and an undetermined
dioecious plant, 30 feet in height, with various other trees,
form a wood abounding in Mosses, which, however, from
its manner of growth, appears to belong to a vigorous
growth of mountain pines (p. 452), although, even as far as
this, a slender tree-fern, Cyath&a oligocarpa, from 15' to
20' high (extending from 5500' 92000, is met with
(see Annual Report for 1841, p. 449). "But," says
Junghuhn, " we search in vain throughout the island for
another example of such a wood on a mountain-top : all
the mountains, far below this altitude, are either bare,
being covered with lava and crumbled rocks, or overgrown
with grass meadows of Festuca nubigena J. or with social
Casuarinas." Junghuhn estimated the upper forest-limit
on the Tjernai volcano (p. 235) at 7000'; it is formed by
Podocarpus imbricata Bl., and is immediately succeeded
by the subalpine shrubs (see preceding Ann. Report),
and this appears to be the general manner in which the
forests are distributed throughout the island. The true
climatic arboreal limit of Java, which is only attained on
the Pang-Gerango, and is here indicated by the mountain-
pine formation of the wood on the mountain-top, is thus
situated several thousands of feet higher than the apparent

376 GEOGRAPHICAL BOTANY.

one, which is merely produced by local conditions of soil,
and thus Junghuhn, by his ascent of this mountain, has
thrown some light upon an anomaly which has hitherto
been almost inexplicable, viz. that the tree-limit in Java
is so much lower than in the Himalaya, and that in
general subalpine Ericaceous shrubs, with the northern
alpine genera (e. g. Ranunculus, Viola, and Gentiand),
descend there to an equally low level of 7 8000'. Yet
the difficulty in explaining these deviations is not, in fact,
completely removed by these observations, but merely
confined within narrower limits ; for although Pang-Ge-
rango teaches us, that at 9200' the most luxuriant woods
still imitate the crooked stems of the mountain -pine, yet
we find in India forests of tall fir trees at a level of more
than 10,000'.

At the foot of the mud-volcano Galungung, Junghuhn
describes the occurrence of almost impenetrable rush-
formations, the marshy surface being thickly covered with
SaccJiarmn Klaga, 15' in height, around which an Equi-
setum and Epidendra are coiled. Above these marshes,
on the slope of the mountain, the forest of Urticeae and
Magnoliaceae commences, including all those accessory
components which render the attempt to describe tropical
forests apparently impossible, even although we should
not aspire to represent its copiousness by words and
expressions, but merely to seize the distinctions in its
mode of development and the conditions under which it
occurs.

Just as the Rosamala-forests in the west of Java deter-
mine the physiognomy of the mountains, when covered
by them, so in the eastern portion of the island do the
forest-regions of Casuarina equisetifolia, which, however,
are not met with below a level of 4000'; hence, although
they ascend higher than other forms of trees, they are
confined to the more limited space on individual elevated
points. No trace of Casuarinte is found west of Merapi,
a mountain from which they are almost extirpated, whilst
they do not appear to be absent from any of the moun-
tain-tops which ascend on the east of it (p. 372).

GEOGRAPHICAL BOTANY. 377

Junghuhn gives the following statements regarding the
altitudinal limits of some branches of cultivation in Java.
Coffee might probably be cultivated as far as a level of
5000', but at present the plantations do not usually
extend beyond 3000' or 4000' (p. 234). Artocarpus
integrifolia and Arenga saccharifera 3000', Duris zi-
betUnw-r-WW (p. 419).

Kittlitz gives two landscapes of districts in Manilla
(Plates XXIII, XXIV), which, like all the others, are
extremely characteristic, but deficient in sufficient bota-
nical elucidation. Montagne has described the Algce of
the Philippine Islands from Gumming' s collections (Lond.
Journ. of Bot, 1844, pp. 658-6:2).

Ill AFRICA.

Of the botanical investigations of the French in Algeria
but few notices have yet been published. Durieu met
with extensive forests of cedar at Blidah, on the Lesser
Atlas (Comptes rendus, vol. v, p. 18). As far as a level
of from 7 800m. the mountain-slope was inhabited,
and the soil cultivated ; the oak then began to be inter-
mixed with the fruit trees, and soon after single majestic
cedars, 40 meters in height, were seen. But it was only
on the southern declivity that the traveller met with con-
nected forests of this tree, which are cut down annually
by the inhabitants ; they do not, however, appear to be
destroyed as at Mount Lebanon, but are apparently readily
reproduced. At Mascara, Durieu found Cattitris quadri-
valvis common, and increasing in frequency thence to-
wards the south (Comptes rendus, vol. v, p. 19). Bory
de St. Vincent has described some new species of Isoetes,
partly living upon a dry soil, from Algeria (1. c. vol. xviii).

We may now recur to Russeger's travels (Annual
Report for 1842), since his work has proceeded to a
considerable extent, and commenced the illustration of

378 GEOGRAPHICAL BOTANY.

the conditions of the climate and soil in a more tangible
and definite style than was the case in the first volume,
which treated of the East (Reisen in Europa, Asien, und
Africa, Bd. 2. Stuttgard, 1843-45. In 1844, appeared
the first part of this volume, including Egypt and Nubia,
and the first number of the second part, containing
Eastern Soudan). The climate of Lower Egypt, as far as
Cairo, is that of the Mediterranean a wet winter (ii,
p. 263) and a serene summer. In Cairo we find the
rainless zone of the north of Africa. At Cairo, according
to the quinquennial average of Clot Bey, there are twelve
rainy days in the year, with 0'034m. of rain. The ab-
sence of rain, both in Upper Egypt (Cairo to Nubia) and
in the Sahara, depends upon constant north winds ; hence
Egypt is climatically a part of the Sahara.

The swollen state of the Nile, produced by the tropical
rainy season, lasts from June to the end of September
(i, p. 229). The months of October and November form
the period at which the Cerealia are sown in those tracts
of ground which are artificially flooded by canals ; the
harvests occur in February and March. Here, accord-
ing to the kind of grain, a second crop may be sown in
April, and reaped immediately before the irrigation. In
other fields, the crop cannot be sown until December or
January, and only once reaped, in May.

Sketch of the most important Branches of Cultivation, arranged according
to the usual period at which the Crops are Sown and Reaped.

SOWN. HARVEST.

January. Beans (Cerealia). Sugar-cane.

February. Rice, maize. Barley, melons.

March. Cotton. Cerealia, maize.

April. (Cerealia.)

May. Tigs, dates, grapes (Cerealia).

June. Beans (Cerealia).

July. Cotton.

September. Oranges, olives, rice.

October. Cerealia. Rice.

November. Cerealia. Maize.

December. (Cerealia.)

GEOGRAPHICAL BOTANY. 379

In the rainless zone of the north of Africa, in conse-
quence of the duration of the polar currents, the great
diurnal differences of temperature allow of the formation
of dew to a slight extent ; it occurs very copiously in the
lower valley of the Nile, is formed in Upper Egypt, and
appears also to fertilize the oases. Russeger, however,
did not meet with any dew in the Desert of Nubia, but
in that of Lybia it is common (ii, p. 253). The oases
lying to the west of Egypt, according to Russeger, gene-
rally obtain their soil-water from the Nile, which flows
laterally to them over beds of clay (p. 271). Thus they
form a valley which is filled with springs, excavated
below the level of the Nile, and parallel to this river.
The other oases of the Sahara appear to be produced
merely by the formation of dews. Borgu, Darfur, and
Kordorfan, however, in this sense are not oases, but
savannahs, situated within the rainy climate (p. 283).

The tropical rains extend in most years to, at the
most, 18 N. lat. (i, p. 224), i. e. two degrees north of
Chartum, the point at which the two arms of the Nile
become confluent. The heavy rains fall there in the
summer, and correspond to the south winds which blow
at this time, and which prevail below 15 N. lat. from
April to September, and alternate every six months with
the north winds. The northern border of this monsoon-
zone, which in the south of the Desert or Soudan pro-
duces savannahs, is not accurately determined. A short
rainy season may occasionally occur beyond 18 N. lat.,
when the south winds blow as far as this part. How-
ever, the dry chamsin of the Desert, which blows from
the same direction, and which Russeger regards as a
local and electrical phenomenon, must not be confounded
with these general south winds which bring rain. Even
between 16 and 18 N. lat. the rainy period is irregular,
and in many seasons abbreviated : at Chartum it lasts
five months. Russeger assumes the following mean
values as marking the north border of the tropical rainy
zone throughout Africa : 21 N. lat. at the Red Sea,

380 GEOGRAPHICAL BOTANY.

18 at the Nile, 16 north of Tschad (according to
Denham), and 20 in Senegambia (ii, p. 546). He forms
a law on the great diurnal differences of temperature be-
tween the night and day, even within the rainy zone,
which, if generally confirmed, would constitute a charac-
teristic peculiarity of tropical Africa.

The whole of Nubia, as far south as 18 N. lat., ex-
cept the valley of the Nile and the coast, consists, like
Egypt, of rocky and sandy deserts. The heights here
extend scarcely 1000' above the plain, on the coast only
ascending to 4000', and in Dschebel Olba, according to
Wellsted, to 8000'. The coast of the Red Sea is not
free from rain ; but on the Nubian side, the summer
rains produced by the south-west monsoon extend almost
as far as the latitude at which the tropical winter rains
(as in Lower Egypt) commence. Suakim is situated on
the northern border of the full rainy season (19 N. lat.);
here, however, it occurs six weeks later (middle of July)
than below 17, and the summer rains are proportionately
retarded and abbreviated as far as 21 N. lat., from which
latitude northwards the winter rains commence. Al-
though the upper portion of the sea north of Suakim is
set in motion throughout the entire year by north winds,
yet the African coast of the Arabian Sea is never free
throughout the entire year from humid currents of air.
This explains how it is that the entire coast line of Nubia
is furnished with willows and other trees, whilst the inner
country does not contain even oases. During the journey
through the Desert, from Korosco to El Muchaireff,
which occupied fifty hours^ and is usually made to avoid
the great bend of the Nile, Russeger only once met with
brackish water, and that was in the middle of the
journey.

The Nile leaves the zone of tropical rain at the
influx of the Atbara, and again comes in contact with it
by its bend at Dongola for a short distance. South of
the mouth of the Atbara, savannahs begin to alternate
with tropical forests, and this is the case throughout

GEOGRAPHICAL BOTANY. 381

the whole of Soudan : no more deserts are met with,
except where the soil is rocky ; they gradually pass into
savannahs (ii, p. 525). The savannahs, during the
rainy season, are overgrown with thick grass ; in the other
months they resemble a dry stubble-field. The forests
consist of Mimosa, and are crowded along the banks of
the stream, as in Guiana. Near the rivers the rain dis-
trict also extends further north ; hence, at a considerable
distance from them, even beyond the 18th degree, the
creeks of the desert encroach upon the savannah.

Throughout the entire district of the Nile, at least as far
as the 10th degree south, there are no terrace-like eleva-
tions of the soil west of Abyssinia, only immense plains.
The terraces of Sennaar, Eazokl, &c., are geographical
over-estimates (ii, p. 539). According to Russeger's
barometrical measurements, the following places are
situated at the annexed altitudes above the Mediterra-
nean : Assuan (Syene), 342', Par.; Korosko, 450';
Abuhammed, 963'; El Muchaireff, 1331'; Chartum,
1431'; Torra, on the White Nile, 1595; Eleis (13),
1667'; and the capital of Kordofan, El Obeehd, 2018'.
Russeger found the northern limit to the occurrence of
Adansonia, in the savannahs of Kordofan, to exist below
14 N. lat.

On the coast of Adel, on the road from Tajura to the
foot of the mountains of South Abyssinia, according to
Harris's report of his travels (The Highlands of Ethiopia.
London, 1844, vol. i, p. 412), the entire country was
desert, and almost dried up in June, i. e. before the com-
mencement of the rainy season, and the soil entirely un-
cultivated. When the heavy rains commenced it was
stormy and unhealthy ; one of the most uninhabitable parts
of Africa, The flora was uncommonly poor ; the woody
plants consisted of shrubs of Mimosa arid Gadaba Indica,
one of the Capparidacese ; subsequently isolated Palms,
Cucifera Thebaica, and below 11 N. lat. Phoenix were
met with. The only other plants found at the end of the
dry season were a few Capparidacese and Malvaceae ; and

382 GEOGRAPHICAL BOTANY.

of other botanical groups of the steppes, single forms only,
as Stapelia, Pergularia, and some succulent Euphorbias ;
but at the river Hawasch the vegetation became social,
by the formation of thickets of Tamarix or Balsamo-
dendron Myrrha, with single Capparidaceous trees
(i, p. 416). At the foot of the high mountains of
Abyssinia, Aloe Socotrina was also met with, and soon
after Tamarix Indica, with which the desert steppe was
overgrown.

Harris read a paper on Balsamodendron Myrrlia before
the Linnsean Society (Ann. Nat. Hist. xiii,p. 220). This
important shrub is called by the Danakil tribes, who
inhabit the coast of A del, Kurbeta. Myrrh (Hofali) is
the milky juice which escapes from wounds made in it,
dried in the air ; it is usually collected in January, the
period at which the buds unfold, and in March, when
the seeds are ripe. Balsamodendron Opobalsamum grows
on the opposite Arabian coast, at Cape Adem. The
Frankincense trees of the mountains of Cape Guardafui,
have not been botanically determined.

Harris's botanical reports upon Shoa are very un-
satisfactory (The Highlands, &c. ii, pp. 395 et seq.) The
pine of North Abyssinia is replaced in Shoa by the
Det, a Juniperus, 160' in height, with a stem from
4' to 5' in diameter, and with the growth of a cypress.
Forest trees are also mentioned : Taxus (Sigba), Ficus
(Schoala), and F. Sycomorm (Work a) ; moreover,
Riippel's Lobeliaceous tree, Rhyncliopetalum montanum
(Jibera), is common at Aukober, the stem of which is 15'
in height, and bears a crown of large leaves. Shrubs :
an Erica (Asta) and Polygonum frutescens (Umboatoo)
distributed generally. Celastrus edulis (Choat) is cul-
tivated commonly, and resembles tea in its action and
taste (ii, p. 423).

Harris's meteorological observations, which were made
from August to December in 1841, and from January to
July in 1842, in Aukober, the capital of Shoa, are of
more importance. This place is situated below 9 35'

GEOGRAPHICAL BOTANY.

383

N. lat., 8200' above the level of the sea, and upon an
open, cultivated flat. The climatic values are as follow :

Mean Temperature.

Number of
Rainy Days.

Quarter of the Wind.

January

11 1 C.

East

February

12 5

East and South

March

East

April

12 9

14 (storms ?)

East

May .

15 4

East

June .

16 7

East

July .
August
September

14 5
13 2
13

H 1 rainy
13 | season

Changeable
Changeable
North and East

October .

North and East

November

North and East

December

East

Mean temperature, 13 1 C. ; Maximum, 20 6 ; Minimum, 5 C.

In Koolo (4 N. lat.), south of Enarea, on the confines
of the pigmy Doco-negro tribes, according to the reports
of the natives, the rainy season lasts from May to February
with but slight intermission (iii, p. 64). To the north-
west of this part, below the 5th degree of north latitude,
the country of Susa is situated, high up on the prolon-
gation of the Abyssinian mountains, and there, as at
Shoa, the rainy season lasts only three months ; but it
must be colder there, for the mountains appeared to reach
the sky, and were covered with perpetual snow. This is
the district in which Bruce supposed the White Nile to
arise.

Hochstetter has described some new Grasses found in
Nubia and Abyssinia, from Kotschy and Schimper's
herbarium, and after making some critical remarks upon
the results obtained by Raffeneau, Endlicher, and himself
in this branch of the Flora, proposes the following new
African genera (Ratisbon flora, 1844): Chasmanthera,
a Menispermaceous plant from Abyssinia; Paulo-Wil-
helmia (Dombeyaceae), from Nubia ; also from Abyssinia,
the Umbelliferse, Agrochoris, Haplosciadium, and Gymno-
sciadium ; Discopodium (Solanacese) ; Hymenostigma, and

384 GEOGRAPHICAL BOTANY.

Acidanthera (Iridaceae) ; and Clinostylis (Liliaceae). Fre-
senius has written notices of some Abyssinian plants
(Bot. Zeitung, 1844, pp. 353-7). Fenzl has announced
a work upon Kotschy's collections from Africa, and in it
has enumerated a series of new forms, but without giving
any descriptions (Ratisbon Flora, 1844, pp. 309-12).

A valuable notice of the plants collected by Krauss in
the most southern regions of the Cape Colony, and in
Natal, with a report on his travels, and a botanico-
geographical introduction, has been published by the
author (Ratisbon Flora, 1844, p. 46). He accurately
describes the large elevated forests, which, are limited
to a small area, considering the extent of the whole
colony, and extend along the south coast between Gauritz
and the Kromme river and the foot of the Onteniqua
mountains. According to his account, Drege's represen-
tation, contained in E. Meyer's work, of the generally
poor character of the forests of the Cape, is not perfectly
correct. At least in this district there exists a quantity
of timber collected into woods, which Krauss characterises
as impenetrable thickets. He mentions some gigantic
stems of Podocarpus, which four men cannot span ; more-
over Crocoxylon excelsum (Safranhout), Ocotea bullata
(Stinkhout), Curtisea faginea (Hassagaihout), and Elce-
dendron Capense trees., the large, densely leafy crowns
of which are elevated far above the thicket beneath, and
covered with numerous climbing plants. Underwood,
e. g. Burchellia, Gardenia, CantJiium, Plectronia, Tecoma,
Grcwia, Sparmannia, and Rubus. Lianes : Cissus, Cle-
matis, Cynoctonum, and Secamone ; Ferns in the deeply-
shaded parts. After a tedious ascent, and laborious
struggle through the chaos of bushes, is finally reached a
thin wood, the trees become smaller and smaller, and
their limit is soon attained. They are succeeded by
shrubs of Synantheraceae, Thymeleaceae, Bruniaceae,
Proteaceae, and Ericaceae.

Krauss confirms the statement that the river Camtos
constitutes a distinct limit of vegetation. This stream

GEOGRAPHICAL BOTANY. 385

might form the boundary between the flora of the Cape,
and of the CafTre country ; for here certain types of tropical
Natal commence, whilst the Proteacese, Ericaceae, Sela-
ginese, &c., diminish. The shrub-formations in Algoa
Bay are taller and thicker than in the western districts ;
they serve as places of concealment for large Pachyderinata.
Characteristic . forms of plants : Celastrinea3, Euphorbia
Canariensis, Strelitza, Zamia, Tamus, Pelargonium, &c.
This remarkable difference between the eastern and
western provinces of the Cape Colony, which Bunbury
also (London Journal of Botany, 1844, pp. 230-63) men-
tions and enlarges upon more in detail, is by no means
to be so simply explained as the tropical peculiarities
of the flora of Natal. At Graham's Town in Albany,
Bunbury only found 13 plants in the extensive surround-
ing country, and these occur also at the Cape. Ericaceas
and ProteaceaB are rare, arborescent Euphorbias common,
and the Restiacese replaced by Grasses. Extending along
the Great Fish River, we find the wildest thickets of shrubs
with arborescent Euphorbias, Strelilzia, and Zamia horrida;
these are more impenetrable, and from the presence of
spinous trees, more inaccessible than the natural Brazilian
forests : they merely form the abode of large Pachydermata
and border-robbers of the Caffre race. Tropical families
of plants, single species of which only occur at the Cape,
become numerous in Albany, especially Acanthacese,
Apocynese, Bignoniacese, Rubiacese, and Capparidaceae.
These and other similar facts evidently indicate an ap-
proximation to the flora of Natal, although by no means
to the extent these two authors suppose, viz. that the
vegetation of Albany and Natal gradually run into each
other. As long as the intermediate districts of the Caffre
country are so little known, this must remain hypothetical,
but is rendered extremely improbable by climatic laws.
A resemblance of certain families and forms is by no
means a resemblance of species and their combination
into formations. But the increase of tropical forms in
Albany is even considerably more mysterious than the

386 GEOGRAPHICAL BOTANY.

contrast between Albany and the west of the colony. In
explaining the latter, we must bear in mind the narrow
district throughout which the Cape plants are distributed,
the tropical forms appear to indicate climatic influences
which do not exist ; for Albany is, if anything, unusually
dry in comparison with the other regions of the colony,
except the district of the Gareep. Rain, says Bunbury
(p. 247), is rare and uncertain ; when precipitations do
occur, it is only the case during south or south-east sea
winds. The climate is indeed considered very healthy,
yet it is subjected to great and sudden changes of tem-
perature, with stormy and dry winds from the west and
north. Hence Albany does not exhibit any trace of that
periodical rainy season, which at Port Natal, as the most
southern point (30 S. lat.) of the regular tropical seasons,
gives rise to the trade- wind character of the flora, and yet,
in so dry a climate, the mode of formation of the plants
is more similar to that of the trade-wind flora, than at the
Cape, where in the winter regular precipitations occur
almost as in the south of Europe. We must, therefore,
in Albany, admit the occurrence of one of those botanico-
geographical facts, where even a tropical constituent of
the vegetation appears to be dependent not only upon
climatic conditions, but historical or geological events.

According to Krauss, Natal is well watered by nume-
rous rivers, which arise in the coast-chain Quathlamba ;
these mountains are nearly 10,000' high, and run through
the coast-country of the new colony in every direction.
The vegetation springs up in September, and during the
months of October, November, and December, corre-
sponding with the atmospheric precipitations, attains the
greatest splendour. During this moist season, the ther-
mometer varies between 19 and 31 c. Vegetable life
is suddenly arrested as early as January, the grass plains
appearing dark yellow, and the forests flowerless and
uniformly green. Rain seldom falls from January to
March ; the air during this period is hot and oppressive,
and the temperature between 26 and 32 5 c. The

GEOGRAPHICAL BOTANY. 387

same appears to be the case with the two following
months, which Krauss did not spend in Natal ; July and
August are fine, the days hot (as high as 31), but cool
in the morning and evening ; the thermometer, however,
seldom falling so low as 15 c. A changeable, windy,
and disagreeable period begins in September the pre-
cursor "of the rain. From these statements, the course of
the seasons is the same as in the East Indies, except that
the rainy season of three months occurs during the spring
in the southern hemisphere, i. e. three months later than
in the former.

Sketch of the predominating botanical formations :

1. Coast or forest region.

a. Forests of Rhizophoreae in the mud between the
ebb and flow of the tide (Mengerhout of the colonists).
Brugmem Gymnorrhiza } Rkizophora mucronata, Avicennia
tomentosa.

b. Dense, tropical, mixed forests, which can only be
traversed by the paths formed by the elephants and
buffaloes. Among the trees several belong to the new
genera published by Hochstetter, with Ficus, Tabernce-
montana, Zygia, Milletia, Phoenix reclinata, &c. Under-
wood, Lianes, and the other components of tropical forests
are copiously developed.

c. Grass plains with various shrubs. Musa.

2. Hilly region, with beautiful pasture land, consti-
tuting the flower of the colony. The woods consist of
Acacias. The Aloe and tall-stemmed Euphorbias re-
semble those in Karro. The highly nutritious grass,
which consists principally of Andropogineae, contains nu-
merous shrubs, especially tropical forms of Leguminosae,
Scrophulariaceae, Labiatae, Acanthaceae, and GnidiaKraus-
siana.

3. Mountainous region. The above-mentioned exten-
sive grass plains are succeeded upwards by a woody belt
of Podocarpus, with numerous Ferns, and above this
mountain-meadows of Cyperaceae, with Orchidaceae, Ixia,
Hypoxis, and Watsoma, are distributed. The largest

388 GEOGRAPHICAL BOTANY.

number of the representatives of the botanical forms of
the Cape occur in this region ; but hitherto only two Pro-
teaceae, one Aspaldthw, two Geraniaceae, one Muraltia,
one Maternia > and one Sarosma t have been found in
Natal, and not a single species of Erica, Phylica,
Selayo. Oxalis, Zygophylleae, &c.

The summary of Krauss's Herbaria contains the diag-
noses of several new species from Natal, and some from
the Cape colony, published under the authority of those
naturalists who have worked out the collections for the
traveller. Among them the following new genera are
proposed : By Bischoff, Splicerotliylax (Podostorneae) ; by
Meissner, Bunburya (Rubiaceae) ; by C. H. Schultz, Mo-
nopappus (Helichryseae) ; and Antrospermum (Arctotideae).
Kunze has described some new Ferns from the Cape and
Natal (Lmnsea, 1844, pp. 113-24).

Bojer has continued his descriptions of new species of
plants from the Mauritius and Madagascar (Troisieme
Rapport de la Soc. de St. Maurice) ; on this occasion they
refer to the Anonaceae, Menispermaceae, Capparidaceae,
and Leguminosae. Gardner has made a brief report upon
some excursions in the Mauritius (London Journal of Bot.,
1844, pp. 481-85).

IV. ISLANDS OF THE ATLANTIC OCEAN.

Seubert has published a copious Flora of the Azores,
in which his former memoir, which has been noticed in
this work, is satisfactorily carried out, and brought to
systematic perfection (Flora Azorica, Bonnae, 1844, 4to).
Of about 400 plants from the Azores, upon which his
observations are made, fifty sp. are endemic, twenty-
three sp. belong also to the Canarian Archipelago, five
sp. to the continent of Africa, and six sp. to that
of America; the remainder occur also in Europe. Of
the endemic species, seven are Synantheraceae, as many
Cyperaceae, and five Graminaceae. Immediately after the

GEOGRAPHICAL BOTANY. 389

appearance of the Flora Azorica, Watson published a list
of the plants which he collected in the Azores (Lond.
Journ. of Botany, 1844, pp. 582-617); and thus increased
the number of the Phanerogamia of these islands, which
have as yet been made known, to about sixty species.
As the plants belonging to the south of Europe found
there are of less interest, we shall confine ourselves to
his contributions to our knowledge of the endemic flora.
He has admitted the following of Seubert's species into
this category : Plantago Azorica Hochst. as a variety of
P. lanceolata, and Juncus lucidus Hochst. as a synonyme
of J. tenuis W. ; also Luzula purpureo-splendens S., ac-
cording to an older syn. L. purpurea Watson; and
Bettis Azorica as a distinct genus, denominated Seubertia.
Lastly, he has described five new endemic forms : Hype-
ricum decipiens (H. perforatum S. ?), Petroselinum tri-
foliatum y Campanula Vidalii, Myosotis Azorica, and
Euphrasia Azorica (E. grandiflora Hochst.?) Vaccinium
cylmdraceum Sm. appears different to him from V.
Maderense Lk. ; but Erica Azorica Hochst. only a var.
of E. Scoparia. The following may be mentioned as
interesting discoveries of the plants of Madeira, and other
adjacent floras in the Azores : Melanoselinum decipiens
Hoffm., Tolpis macrorrhiza D. C., Mirabilis divaricata
Lour., and Per sea Tndica Spr.

Seventy-five parts of the work of Webb and Berthollet
on the Canary Islands are out. They carry the systematic
part as far as the Synantheraceae.

Reid has communicated some reports upon the cedar of
the Bermuda Archipelago (Lond. Journ. of Bot., 1844,
p. 266, and 1843, p. 1). The inhabitants erroneously
consider this Coniferous plant (Juniperus Bermudiana)
to be the same as the Virginian cedar (Juniperus Fir-
giniand}. Even the climate of these islands is very
different from that of the opposite coasts of the American
continent, as water never freezes in the Bermudas. The
most magnificent oranges are produced there, being pro-
tected from the winds of the Atlantic by the large forests

390 GEOGRAPHICAL BOTANY.

of these cedars, which cover all the uncultivated regions.
This tree is also called the pencil cedar, although the
wood does not appear to be used at present in the manu-
facture of lead-pencils in England. It is much prized
for ship-building. Reid thinks that the Bermuda cedar
does not occur in the hot climate of the West Indies, but
it is very common on the mountains of Jamaica.

V. AMERICA.

The plants collected by Simpson and Dease, in their
voyage of discovery on the arctic coast of America, have
been named by Sir W. Hooker (Narrative of the Disco-
veries on the North Coast of America, by T. Simpson.
London, 1843, Svo, Appendix). These plants had, how-
ever, been previously found in Eranklin's travels in the
same region, and admitted into Hooker's Flora of British
America, with the single exception of Salix nivalis
Hooker, which was discovered by Drummond on the
Rocky Mountains, and occurs also on the coast below 71
N. lat., west of Mackenzie.

A coast-landscape of Unalaschka, by Kittlitz (pi. IV),
represents luxuriant meadows, in which various subalpine
shrubs, forming most luxuriant thickets of plants, are
intermixed with strong turf composed of Cyperaceae :
amongst the former are Aconitum, Heracleum, JEpilobium,
and especially Lupinus. The dwarf shrubs, also, of the
alpine region, Salices and Rhododendron Kamtschaticum,
extend on these islands, which are situated beyond the
tree-limit, into the vicinity of the sea. Two views of the
island of Sitcha, the forests of which they represent (pi.
II and III), may serve as contrasts. They give a dis-
tinct representation of the mixed foliage of the Canadian
larch (Pinus Canadensis) and a species of pine (P. Mer-
tensiana), the growth of Panax horridum, the palmate
auricled leaves of which are sometimes crowded together
upon a turfy kind of brushwood, at others upon shrubby

GEOGRAPHICAL BOTANY. 391

stems, consisting of the Vaccinia and Rubi, forming the
underwood, and other botanical forms, with which Bon-
gard's sketch has made us acquainted.

After an interval of two years, the third part of the
second volume of Torrey and Asa Gray's 'Mora of
North America/ containing the completion of the Synan-
theraceee, has appeared. A. Braun has described the
Equiseta and Charte of North America (Silliman's Journal
of Science, vol. xlvi). Mac Nab read before the Edin-
burgh Botanical Society a botanical journal, which he had
kept at Hudson (Ann. Nat. Hist., xiv, pp. 223-25).

Asa Gray has continued the report upon his botanical
journey in the south of Alleghany (London Journ. of
Bot. 1844, pp. 230-42). On the summit of the Roan,
in Tennessee, the altitude of which is 6000', Rhododen-
dron Catawbiense forms a fertile subalpine shrubby forma-
tion, the turf of which consists of Carex Pennsylvanica
and other species of this genus, with Aira flexuosa and
Juncus tennis. Beneath the shrubs, Lilium, Veratrum,
Potentilla, Geum, some Ranuneulacese, Umbelliferse, Saxi-
frageae, and Solidago, with Rudbeckia, Liatris, &c. ? are
mentioned. The remaining woody plants, in addition to
the Rhododorese and Rosaceae, mentioned in the Annual
Report for 1842, consisted of Pyrus arbutifolia, Cratcegm
pnnctata, Ribes rotundifolium, Diervilla trifida, Vaccinium
Constablcei n. sp., and Alnus crispa. Pinus Fraseri is
the tree which extends to the greatest altitude ; it occurs
near the summit in a dwarf and crooked form. At the
end, A. Gray describes the new genus Shortia (yala-
cifolid) from specimens in fruit in the herbarium of
Michaux, who discovered it on the mountains of Carolina.
It has not since been found, and its flowers are unknown.
This remarkable plant unites the habit of Pyrola uniflora
with the leaves of Galax. Nuttall described another
genus (Simmondia) from S. Diego, in Upper California,
as a new type of the Garryacese (1. c. p. 400, t. 16).
The collections of Hinds (Ann. Rep. for 1842) have
afforded the matter for an important systematic illustrated

392 GEOGRAPHICAL BOTANY.

work, which Bent-ham is working out, and the traveller
elucidating by botanico-geographical remarks (The Botany
of the Voyage of H.M.S. Sulphur. Edited and superin-
tended by R. Brinsley Hinds. The botanical descriptions
by G. Bentham. London, 1844, 4to). Five parts are now
published. The representation of the character of the
vegetation of California given in this work has a decided
preference over the earlier ones, upon which we have pre-
viously reported. The flora of California resolves itself
into two districts, a northern one, extending from the
Columbia river to S. Diego (33 N. lat.), and a southern
one, from here to the vicinity of the tropic, where the
tropical forms of plants commence : the former corresponds
to about the limits of Upper, and the latter to those of
Lower California. South of Columbia (46), where the
forests of Abies terminate, the deciduous forests gradually
continue to disappear: above S. Francisco (38), there
are no large forests, and altogether but few trees. In
sailing, in Upper California, up the S. Francisco from the
coast, a broad alluvial plain presents itself; it is open,
and here and there sparingly wooded with oak trees like
a natural park : the river flows through it, and floods it
in the moist season. Bentham determined the trees to
be, Quercus agrifolia and Hindsii and Oreodapkne cali-
fornica ; Frawinus latifolia and ^Esculus calif ornica are
also found there, and Salices, with Plat anus calif ornica,
grow upon the banks of the river. At S. Pedro, the
flora of Lower California prevails, and extends as far as
the Magdalene Bay (24 38'), where the most northern
mangrove forests are found. Between these two points,
the soil at different landing-places was either covered with
low shrubs, which frequently fill the air with agreeable
odours, or (in October and November) bare, like the
steppes, and ornamented between the isolated portions of
underwood with herbaceous plants with very beautiful
flowers. Here the Synantheracea3 predominate, in the most
varied forms and colours ; in fact, they constitute more
than the fourth part of Hinds's collection. Next to these,

GEOGRAPHICAL BOTANY. 393

the Euphorbiaceae, Polygonaceae, and Onagrariae are more
abundant than the remaining families : the entire Cali-
fornian herbarium, however, only includes about 200 sp.
The arid and frequently sandy soil is physiognomically
characterised by different Cacti, 2 of which are distributed
exactly as far as S. Pedro, and accurately define the
extent of the flora. At Magdalene Bay, other tropical
forms also begin to appear with the mangrove forests in
considerable numbers, which are mixed in the text with
the steppe- plants of Lower California, but which must of
course be distinguished geographically from them. The
Euphorbia-shrubs only are common to both the districts
of the peninsula; nevertheless, the species which pre-
dominate within and without the tropic are different.
Magdalene Bay appears to form a well-defined floral limit
northwards. Together with Cape Lucas, it yielded one
half of the plants contained in Hinds's Californian herba-
rium. But whether this tropical southern point of the
peninsula forms a distinct and third botanical district, or
should be considered as belonging to that of the western
coast of Mexico, remains at present undecided, inasmuch
as most of the plants collected here have not yet been
described. The following are the families of the latter
collection which contain the largest number of species :
Synantheraceae Q), Euphorbiaceae Q), Leguminosae (],
Graminaceae, Solanacese, Malvaceae, and Nyctagineae.
New genera from California, by Bentham : Stegnosperma
(Phytolaccaceae),/$'m^^te andEremocarpus (Euphorbia-
ceae), Helogyne, Perityle, Coreocarpus, Acoma, Amauria
(Synantheracete), Eriodictyum (Hydroleaceae). F. D.
Bennett in a short time collected some 70 sp. on the
tropical south point of California ; they have not yet been
made known (Narrative of a Whaling Voyage. London,
1840, vol. ii, p. 18). He saw there columnar Cacti,
from 15' 20' in height, and speaks of the luxuriance
of the forests and of numerous succulent and bulbous
plants.

Martens and Galeotti have continued their papers on

394 GEOGRAPHICAL BOTANY.

the flora of Mexico (Bullet, de FAcad. de Bruxelles,
1844, vol. xi, part 2, pp. 61, 185, 319; 1845, vol. xii,
p. 129) : they contain 74 Labiate, with the new genus
Dekinia, 39 Verbenaceee, 9 Cordiacese, 30 Boraginacese,
and 63 Solanaceae. The Ferns (170 sp.) and Lycopo-
diacea? (12 sp.) are treated of by them in detail (Memoires
de FAcad. de Bruxelles, 1842), and are illustrated by
copper-plates. Kunze has described the Ferns and the
allied families collected by Leibold in Mexico (128 sp.)
(Linnaea, 1844, pp. 303-52). V. SchlechtendaFs conti-
nuation of his ' Contributions to the Flora of Mexico'
contain the Sapindaceae, a new Dioscoreaceous plant,
and Hydrotania (1. c. pp. 48, 112, 224). Bateman has
published a splendid illustrated work on the Orchidaceae
of Guatemala and Mexico, with 40 plates (Orchidaceae of
Guatemala arid Mexico. London, 1843, imp. fol.)

Galeotti, in his ' Memoir on the Mexican Ferns/ has
also investigated their distribution in the regions which
he has assumed, and commenced a similar work in con-
nexion with Richard, in which the Orchidacese of Mexico,
where, according to Richard's judgment, the forms of
this family are more abundant than in any other country
in the world, are treated monographically, from a collec-
tion of 500 species (i. e. \ of all that are known), and the
geographical distribution of which is given (Comptes ren-
dus, vol. xviii, pp. 497-503) in a preliminary paper. The
regions assumed by Galeotti in these two papers include
the greater part of Mexico, without, however, as was the
case with Liebmann's ' Characteristics of Oribaza,' their
being supported by a sufficient number of special investi-
gations. We cannot judge of the value of Galeotti's
botanical division of the country until, as it is undoubt-
edly his intention to do, a special work is published on
the botanico-geographical relations of all the Mexican
families of plants. The altitudes given do not always
agree with those of Liebmann, nay, in some cases, they
do not agree with each other : how far this is due to
inaccurate observation, and how far to local variation in

GEOGRAPHICAL BOTANY. 395

the limits of the plants, cannot be satisfactorily deter-
mined. In the following sketch of Galeotti's regions,
the local displacements are added within brackets to the
altitudes given.

1. Hot regions. 0' 3000' (25000. Vegetation from
December to May (end of October to June) languid :
most of the trees leafless.

a. Eastern coast with forests of Rhizophoras. Mean
temperature = 95 c.

b. Moist mixed forests, not, however, containing many
Ferns (R. chaude temperee des ravins). Mean tempe-
rature, 25 19 c.

c. Coast forest of the Pacific, 25 19.

2. Temperate regions.

a. Eastern slope. 3000 6000' (5500', 70000. This
region differs from the coast in its great humidity and
evergreen foliage. It contains tree-ferns, Liquid-ambar,
evergreen oaks (a feuilles luisantes), and numerous Orchi-
daceae. Mean temperature = 19 to 15. In Oaxaca,
this region is less distinctly separated from the others :
the pine trees here descend as low as 3000', whilst Myr-
taceae, Melastoma ( , &c., are found even at an elevation
of 7000'. The soil is calcareous, and Galeotti found
there only 21 Ferns, whilst on the volcanic soil of Vera
Cruz, he found 77 species at the same level.

b. Western slope. 3000' (10000 to 6500'. Mean
temperature = 20 to 15. To it a large part of Oaxaca,
of Mechoracan, and Xalisco belong. Tree-ferns are not
found there, and in fact few Ferns, but a large number
of oaks, with numerous Orchidacese growing parasitically
upon the bark, and some Palms.

c. Plateau, and the slopes adjacent to it, mean tem-
perature = 2 0to 15 (21tol8). The internal slopes of
Mexico differ in every case, botanically speaking, from
those situated externally, and inclined towards the two
oceans. Their dry climate, for the most part, excludes
the vegetation of Ferns and Orchidacese. These elevated
surfaces are characterised by the large number of Cacti :

396 GEOGRAPHICAL BOTANY*

spiny Mimosa and unparasitic Bromeliacese are common.
The latter, with Agave, are frequently the only plants
occurring on the calcareous soil; or on other kinds of
mountains, the surface is extensively covered with Pro-
sopis dulcis and Mimosa. Bronnia spinosa is also cha-
racteristic.

3. Regions froides.

a. Eastern slope. The determinations of the altitudes
of the upper stages of vegetation, e. g. at Oribaza, are in
part inaccurate ; thus, according to Liebmann's investiga-
tions, the statement that vegetation ceases at 12,500' or
13,000' is incorrect ; this portion of the sketch is therefore
passed over. This region has yielded 52 Perns, most
of which grow upon limestone, and also numerous Orchi-
dacese (especially between 7500' and 8000').

b. Western slope and high mountains of the plateau.
Botanical characteristics wanting. The upper limit to
vegetation is situated, according to Galeotti, on Popoca-
tapetl, at 11,500', on the Pic of Toluca, at 13,000'.

c. Most elevated surfaces of the plateau. No botanical
characteristics.

The second and larger section of Hinds's andBentham's
work (v. s.), which is, however, not yet perfectly com-
pleted, includes the west coast of America from S. Bias
(21 32' N. lat.), to Guayaquil (2 30' S. lat.) On this
long coast-line the flora is adapted to a moist tropical
climate, and the shore covered with a dense forest ; but
the plants north and south of Panama are not the same.
Nor are the seasons contemporaneous ; the tropical rains
commence at Guayaquil in the beginning of the year;
towards the north, they gradually occur later, so that at
S. Bias they commence at the end of June. They divide
the year into two periods of vegetation; the Bay of Choco
alone forms an exception, for there atmospheric precipi-
tations last from ten to eleven months, producing a vege-
tation which is constantly green, and abounding in flowers.
The forests of Guayaquil appear to be comparatively poor
in forms, because the rainy season there, and with it the

GEOGRAPHICAL BOTANY. 397

luxuriant growth of the plants, in the vicinity of the
Garua, only lasts for a short period. Of the characteristic
tropical forms, some are absent, or rarely found; as
Epiphytes, all the Monocotyledons, and the Ferns. North
of Guayaquil the desert tracts again recur, in which the
coast-stream at Salango (2 S. lat.) clothes a spot of land,
like an island, with tropical trees ; but as soon as the
equator is passed northwards on this coast the vegetation
acquires variety and strength. The Orchidaceae and
other Epiphytes then become more common ; the number
of forest forms rapidly increases in the same proportion
as the duration of the rainy season augments, as far as the
Bay of Choco (3 7 N. lat.), where the vegetation of the
western coast is most copiously developed ; but the solstitial
point is also reached at the same time. In this climate,
the boundary of which is on this side of the equator, but
which is still equatorial, the western coast contains its
only Tree-ferns, and even here the Cacti, the characteristic
plants of the trade- wind flora of America, are absent.
At Panama (9 N. lat.) we again find a proportionate
change of the two tropical seasons, hence no Tree-ferns
nor Scitamineae are met with there, but arborescent Cacti
and other succulent plants. Most of the new species of
the collection described by Bentham are from this south
region of the western trade- wind coast (9N.lat.to3S. lat.)
North of Panama the influx of Mexican types is percep-
tible; Heliantheae become numerous ; the forests of maho-
gany at Realejo are also succeeded above by a region of
Pinus occidentalis, and the oak is found even 15 00' above
Acapulco. 654 species of the rich collection have already
been described in the parts at present published, which
extend from the Polypetalae to the Scrophulariaceae.
Families containing most species: Capparidaceae (10),
Malvaceae (31), Byttneriaceae (11), Sapindaceae (12),
Leguminosae (125), Melastomaceae (23), Rubiaceae (39),
Synantheraceae (95), Apocynaceae (13), Bignoniaceae (17),
Convolvulaceae (39), Boraginaceae (23), Solaneae (25),
and Scrophulariaceae (at present 17). Considering the

398 GEOGRAPHICAL BOTANY.

large number of new species, the number of undescribed
genera is not great : Triplandron (Guttiferae), Pentagonia
(Rubiaceae), Oxypappus (Synantheraceae), Stemmadenia,
3 sp. (Apocynaceae), Diastema (Gesneriaceae), Thinogetum
(Solaneae), and Leptoglossis (Scrophulariaceae).

Purdie, a collector for the Kew Gardens, has reported
upon his travels in the West Indies (Lond. Journ. of
Bot., 1844, pp. 501-33). Among others he ascended
the peak of the Blue Mountains, in Jamaica, where the
forests of the summit consist of Podocarpus coriacea
(Yacca). In other respects these, as also Moritz's Bota-
nical Letters from Cumana and Caracas (Bot. Zeit., 1844,
pp. 173, 195, 431), merely give notices of the plants
collected.

Miquel has continued his Contributions to the Flora
of Guiana (Linnaea, 1844) : some new Capparidaceae,
Sapindaceae, Malpighiaceae, Dilleniaceae, Leguminosae,
Melastomaceae (Hartigia, n. gen.), Memecyleae, Passi-
floreae, Onagrariae, Cucurbitaceae, Loranthaceae, Ru-
biaceae, Convolvulaceae, Cuscutaceae, Bignoniaceae (Cal-
lichlamys = Bign. latifolia Rich.), Avicenni<,Nyci&g\-
naceae, Polygonaceae, Piperaceae (Nemat anther a n. gen.),
Bromeliaceae, Musaceae, ScitamineaB, Hydrocharidaceae,
Commelynaceae, Xyrideae, and Aroideae. Steudel (Ratis-
bon Flora, 1844) : on the Melastomaceae from Surinam,
and various plants in the collections of Hoffmann and
Thappler, which are for sale. Robert Schomburgk
(Lond. Journ. of Bot., 1844, pp. 621-31) : a new Rubi-
aceous plant, and two Lauraceae, from British Guiana.
Berkeley on Stereum hydropliorum (Ann. Nat. Hist., xiv,
p. 3.27).

Richard Schomburgk, who accompanied his brother
during his last travels in British Guiana, has described in
his letters the botanical characters of the explored regions
(Bot. Zeit., 1844-5). We thus obtain an interesting
supplement to the previous work of Robert Schomburgk
on his travels, in which the botanical determination of
the plants was omitted, and which, now that a great part

GEOGRAPHICAL BOTANY. 399

of the previous herbaria have been described, may be added
to the descriptions of the country. The forest at Esse-
quibo, from which Mora excelsa projects to an altitude
of 160', formed the first opportunity for the traveller to
develope his descriptive talent. After having vividly
delineated the crowded growth of the trees, the climbing
plants and the creeping shrubs, which connect the stems
in impenetrable meshes, and the parasites of the fallen
trunks, he dwells upon a point with which we are less
familiar the light of tropical forests. On the ground the
eye would miss the splendour of the flowers of other
regions, and detect only Fungi, Ferns, and decaying
vegetable structures ; for even at noon a subdued light
prevails in the forest, since scarcely anywhere is a portion
of the sky visible through the closely-interlaced branches ;
but although the light is subdued beneath so dense a
covering of foliage, there is more light than in dark pine
forests. V. Kittlitz comes to the same conclusion as to
the remarkable and as yet but little studied question, of
how plants still thrive so well, and their green organs are
able to respire, in shaded parts of the most dense vege-
tation which the crust of the earth anywhere produces
(Vegetations- Ansichten, p. 6). " I was astonished,"
writes he, " to find so much light beneath the noblest
trees, the widely- spread foliage of which scarcely any-
where allowed the sky to be seen. Remaining the same
at the most varied times of the day, it could not be
ascribed to the perpendicular light of noon, but only
to those innumerable undulations of light which, falling
from above through the crowded masses of leaves in every
direction, being reflected from stem to stem and from
branch to branch, finally reach the lower space in the
thicket, and there produce a tone of dull lustre peculiar
to tropical nature. In fact, what would become of that
whole world of plants destined to live in this shade, if
nature had not given the huge masses of foliage, which
produce it, a structure and distribution, which permits it,
although reflected a thousand times, still to reach in suf-

400 GEOGRAPHICAL BOTANY.

ficient power the plants living beneath." This problem
may be expressed more definitely as follows. We have
to explain why the shadow of obscure deciduous forests
in the temperate zone are principally illuminated by
transmitted, and in the tropics by reflected light, and
why the Coniferous forests are poorer in these two lumi-
nous sources, and therefore so frequently deprived of
plants growing in the shade. We first think of the
Mimosa and forms of Palms, of the compound, and
therefore imperfectly shading forms of leaves, which
thus contribute powerfully to the light tone of the tropi-
cal forests. But trees possessing this character form a
part only, not the whole ; for those forms with simple
leaves, as the laurel- and Bombax-type, preponderate, in
variety of form or size of the leaf. And even the form of
the leaves of the Lauracese, which recurs in so many
tropical families, is wanting in that transparent texture
to which the light of the half-shaded parts of the northern
deciduous forests is owing. But Kittlitz has pointed out
another more universal character of the trees of tropical
forests, in the arrangement of the leaves, which appears
intended to complete the former. In climates where cold
or aridity cause the winter-sleep of woody plants, they
develope a very much larger number of small branches,
which usually form a more connected, although on the
whole poorer, stratum of leaves than in the tropics.
This, therefore, throws a deeper shadow upon the
ground, although it is more transparent ; not so deep,
however, as in the Coniferous forests, the crowded
leaves of which are opaque. On the other hand, it is
evident that the uninterrupted heat and moisture of
the equatorial climate also ensure a longer duration of
the first-formed branches, many of which, in the temperate
zone, fall off or remain undeveloped, and must therefore
produce fresh ramifications to allow of the necessary num-
ber of leaves being formed ; these first branches attracting
the currents of sap, continue to grow excentrically, and
hence leave between their uppermost tufts of leaves, i. e.

GEOGRAPHICAL BOTANY.

401

the youngest and softest part, more or less broad intervals.
Under this double condition of the formation and dis-
tribution of the foliage, we may perceive universally in
the latter climate " a certain and wholly peculiar per-
meability" seen only in its simplest and most developed
state in the Palms even in woody plants, which in other
respects but little resemble the latter, and in which the
more copious development of the ramifications of the stem
produces this prevailing character, inasmuch as they
imitate and replace the natural growth of the summit of
Palms. " Large masses of very delicate foliage in this
manner obtain so light an aspect, that they appear as it
were to float in the air ; but, even down to the smallest
Fern upon the soil, everything exhibits a tendency to an
excentric distribution, which does not permit the separate
organs to press upon one another, but by the constant
crossing of lines in every direction, produces spaces for
the transmission of air and light." Here Nature addresses
man like the noblest works of mediaeval architecture,
the pointed arches of which, of Arabian origin, have, it is
supposed, borrowed that openness conjoined with gigantic
masses and infinite variety of form, from two palm-stems
with their penniform leaves in contact.

As the second principal formation of Guiana, R.
Schornburgk describes the vegetation on the banks of the
streams, at the border of the forest, as made generally
known by V. Martius and Poppig, from the north of
Brazil. The underwood surpasses the retreating gigantic
stems ; a belt of Cecropias and bamboos forms the fore-
ground ; herbaceous lianes wind around the trees and
bushes as in a most luxuriant hedge, on the borders of
which beautiful flowering plants augment still more the
most abundant variety.

From Essequibo the travellers went to the tributary
stream, Rupununi, to arrive at the savannahs on the
sea of Amuku, which in these regions cover the ridges of
the land almost down to the water-line, and are only
separated from the rivers by seams of woods from 100'

402 GEOGRAPHICAL BOTANY.

to 200' in breadth. The main mass of the vegetation in
the savannah consists of scabrous, straggling Graminacea3
and Cyperacea3, from 3' to 4' in height, as Pariana cam-
pestris, Cliatospora capitata, Elionurus ciliaris, Sataria
composita, Mariscus Icevis, intermixed with prickly or
arborescent underwood of various kinds, as Curatella
Americana, Byrsonima, Plumieria, Leguminosse, Myrtacese,
some Synantheraceae, and Malvacea3. The marshy places
are denoted by Mauritia flexuosa, with Melastomas,
Scitamineaa, Polygaleas, and Byttneria scabra ; the surface
of the water itself, Pontederia and NymphaeaceaB.

Poppig's illustrated work upon Tropical America is now
completed by the 7 10 decades of the third volume
(Lipsise, 1844, 4to). The 75th to 78th parts of Orbigny's
Travels have appeared. Klotzsch has commenced the publi-
cation of ' Contributions to the Flora of Tropical America,'
from the Museum of Berlin (Linnaaa, 1844), comprising
at present the vascular Cryptogamic plants and the Jun-
germannise, by C. Miiller.

V. Tschudi's zoological work upon Peru contains, in
the introduction, an interesting division of the Peruvian
Andes, according to their climatal conditions and botanical
characters (Untersuchungen uber die Fauna Peruana,
Lief i. St. Gallen, 1844, 4to). The climatal regions of
Peru, the elevated surfaces and valleys of which are pro-
duced by the structure of the two Cordilleras, and are
not dependent upon the polar altitude, are, according to
Tschudi, as follows :

1. Western slope (contains no woods).

a. Coast region (0' 1500'). Mean temperature in
the hot season = 27 c. \ during the garua = 19- 75. A
band of sand, 1620 miles in length, and from 18 to 60
miles in breadth, extending across the rivers, which inter-
sect it many times, subdivides it into two principal forma-
tions ; for the banks of the river form oases of cultivation
in the Peruvian coast-steppe, the barren hilly surfaces of
which are covered with fine quicksand, and are devoid of
springs and, during the dry seasons, of vegetation. This

GEOGRAPHICAL BOTANY. 403

hot and dry season lasts from November until the end of
April. The garua, a thin mist, which is thickest in
August and September, reanimates the steppes from May
to October. It only extends 1400' high vertically in the
atmosphere. As long as this prevails the steppe is verdant,
and sends forth numerous Liliaceous forms into flower.
The south winds generally last throughout the entire year ;
and V. Tschudi considers the formation of the garua as still
unexplained. May they not arise, as winterly precipita-
tions, from an admixture of the lower trade-wind with
the east winds descending from the Andes, and which,
during the summer, are not in a condition to separate
the moisture from the coast trade-wind ?

b. Internal coast-region (1500' 4000'). This com-
prises the fan-shaped expansion of the west valleys of the
Cordilleras, which, at the time of the garua, is affected
with a true rainy season. Mean temp, in the dry season,
= 29 0> 25 ; in the rainy season = 22'75. The vegetation
is not very luxuriant, but the cultivated tracts are extra-
ordinarily productive. The sugar-cane thrives well even
at 3600'. Of fruits, Anona tripetala (Chirimoya) and
Passiflora quadrangular is (Granadilla) are peculiar to this
region.

c. Western Sierra (4000 11,500'), or that slope of
the Cordilleras which is gently inclined below, and steep
above, with its narrow transverse valleys. The air is dry ;
the nights are very cold in summer \ the prevailing wind
is the east. In summer the mean temperature at noon
is = 22'4, at night = 10 ; in winter, the mean diurnal
temperature is = 19. This is the region of the tropical
Cerealia, and that in which the potato thrives readily and
in profusion. Oxalis tuber osa (Oca) commences in it.
The Cacti are among the characteristic plants of this slope,
which contains but little wood.

d. Western Cordillera, comprises the west slope of the
Andes above 11,000', and the east declivity of this
western crest as far downwards as 14,000'. It forms a
wild mountainous tract, containing steep rocky declivities.

404 GEOGRAPHICAL BOTANY.

valleys expanded into small plains, and numerous alpine
lakes, and is bounded by glaciers and perpetual snow.
Catting, ice-cold winds from the east and south-east
prevail constantly. Mean temp, of the day in summer
= +ll-25, of the night = 7-l; in winter, i. e.
during the rainy season, of the day =+ 7'5, of the
night ==+ 2 0< 6. The vegetation extends to 15,500',
and consists of low Cacti and alpine plants.

2. Eastern declivity (two regions containing no forests,
two wooded).

a. Puna region (11, 000' 14,000'), or the large un-
dulating plateau between the two Cordilleras, and which
attains a mean altitude of 12,000'. Sparingly over-
grown flats alternate with extensive marshes, lakes, and
alpine rivulets. Cold west and south-west winds blow
throughout the year, most violently from September to
May, with fearful thunderstorms, which occur almost
daily during these months. Thus the rainy season com-
mences in the opposite half of the year to any one travel-
ling from the coast to the interior. The sky is clear from
May to September, and the nights very cold ; the tem-
perature is altogether very variable ; it frequently varies
within twenty-four hours 22 or 25 degrees; not unfre-
quently, on these cold heights, we suddenly encounter
warm currents of air from the S.S.E., which at times are
only from two to three paces in breadth, while in other
cases as much as several hundred feet, and which rapidly
recur (p. xxiv). Tschudi gives, as approximative mean
values of the temperature : Summer (November to April,
which period is there incorrectly called the winter),
of the night =-f-l-5, of noon=8'75; winter (May
to October, there incorrectly called summer), of the night
= 6 0< 25, of noon=12 0< l. The vegetation of Puna is
poor; Stipa Ichu is abundant; Synantheraceae, Malpi-
ghaceae, Leguminosse, Verbenacese, Scrophulariacae, and
Solaneae are mentioned. Barley does not ripen at
13,050'.

b. Eastern Sierra (11,000' 8000'), consists of broad,

GEOGRAPHICAL BOTANY. 405

open, fluviatile valleys, the most thickly populated in
Peru, and is separated from that of the Puna by rocky
declivities ; rainy season, with frequent hail, from October
to February. During the winter months (also called
summer in the text) dry east winds prevail ; night frosts
set in after the end of the rainy season, and the Cerealia
are harvested. Mean temp, during the rainy season : of
the night = + 5'l, of the day = + 14'l ; during
the winter (March to September), of the night = 4'25,
at noon + 17'l. But great local differences occur in
the hot bottoms of those valleys which are sheltered from
the winds, where fruits of the south of Europe, as peach
trees, thrive, sometimes even at an altitude of more
than 10,000'; the principal cereal grain appears to be
maize. The slope of this region, which, like the former,
is destitute of woods, contains a profusion of Cacti, and
on the banks of the streams only we find woods of 8alix
Humboldtiana, 20' in height ; even European fruit trees
do not thrive when cultivated. In the valleys, however,
this region extends directly into the forest region, from
which it is also separated by a second Puna, i. e. by the
crest of the central Cordillera.

c. Upper forest region or Ceja-region (from Ceja de la
Montague, i. e. the brow of the mountain) (8000' 5500'),
comprises the eastern slope of the internal Cordillera, and
its western slope in the north of Peru, with the longitudinal
valley of Huallaga. It consists of steep rocks and narrow,
wooded mountain-ridges. The climate is humid, cold,
and rough, with prevailing south winds. Towards evening
dense mists are formed, which during the night rest upon
the forests, and which the wind carries away with it from
the morning until the serene evening. These mists ex-
tend downwards as far as 6500', and often resolve them-
selves into very heavy showers. The differences in the
seasons are not mentioned ; the observations upon the
temperature are also incomplete. Low trees and shrubs
covered with mosses commence even at 9500', and
increase in size and strength as we ascend. Cerealia

406 GEOGRAPHICAL BOTANY.

cannot be cultivated in this region, which is not exposed
to the direct rays of the sun ; potatoes grow abundantly.

d. Lower forest region (5500' 2000'), is composed
of immense forests, savannahs, and marshes. Its humidity
is great throughout the year ; for even in the dry season
(May to September) thunderstorms are common. The
true rainy season begins in October, and lasts until
March or April. Mean temp. = 30; when the wind is
in the east, the nocturnal temperature sinks to 18*75.
This region forms the commencement of the primaeval
forests of the Amazon.

Contributions to the Flora of Brazil : Moricarid, Plantes
nouvelles ou rares d'Amerique, livr. 8, Tab. 71-84
(Geneve, 1844, 4to) ; Naudin, Description de Genres nou-
veaux de Melastornacees (Ann. d. Sc. Nat., 1844, ii,
pp. 140-56) : Tulasnea, Brachyandra, Eriocnema, Auyus-
tinea, Stenodon, and Miocarpus ; Fischer and C. A. Meyer,
Aster ostigma, n. g. Aroideae (Bull. Petersb. iii, p. 148).
Miers, Triuris and Peltophyllwn, forming the new family
Triurideae, allied to the Juncagineae (Transact. Linn. Soc.
xix, pp. 77, 155) ; Sir W. Hooker and Wilson, Enume-
ration of the Mosses and Hepaticse collected in Brazil,
by G. Gardner (Lond. Journ. of Bot., 1844, pp. 149-67).
K. Miiller, Enumeration of the Mosses collected by
Gardner in Brazil (Bot. Zeit. 1844, p. 708), gives no
description of the new species, so that the preceding
publication, which is founded on more complete materials,
acquires priority as regards the nomenclature.

Tenore has described a new Aristolochia, from Buenos
Ayres, which he obtained from Bonpland's collection of
seeds, and has taken this opportunity of republishing the
diagnosis of some plants derived from the same source,
and described in his catalogue of seeds (Rendiconto di
Napoli, 1842, pp. 345-8).

Kittlitz's first plate represents the botanical character
of the coast of Valparaiso. It gives a view of one of the
steppes during the dry season, the bare soil of which
appears only to produce Cacti and shrubs with stellate

GEOGRAPHICAL BOTANY. 407

prickles, but in which, during August and September,
the most luxuriant grass plains, with their bulbous plants,
are found. The following are some of the physiognomi-
cally important forms of plants represented in this drawing:
the Caves (Mimosa Cavenict), the dwarf-pine-like Lithi
(Rhus caustica), Cereus Peruvianus, Puretia coarctata,
Synantheraceous shrubs, bamboos, &c.

Miers has proposed two genera of Iridaceae from Chili
Solenomelus (Cruckshankia ej. ol.) and Sympliostemon
(SisyrincJiium odoratissimum Cav.) (Transact. Linn. Soc.
xix, p. 95). Sir W. Hooker has determined the Alerse
tree of the south of Chili, which is used as timber for
building, to be Thuja tetragona (Lond. Journ. Bot., 1844,
p. 144). Berkeley has described an edible Fungus from
Terra del Fuego ; Cyttaria n. gen., near Bulgaria, also
containing a species from Chili (Transact. Linn. Soc., xix,
p. 37).

VI. AUSTRALIA AND SOUTH-SEA ISLANDS.

F. D. Bennett remarks that westerly winds, corre-
sponding to the monsoon, not infrequently extend east-
wards over the Pacific Ocean towards the Society Islands,
and especially in February and March are not infre-
quently taken advantage of, for making voyages in a
south-easterly direction ; consequently in regions which,
in other respects, are completely under the influence of
the south-east trade- wind (Whaling Voyage, i, p. 159).
The botanical communications, which form an appendix
to the account of his voyage, and which treat especially
of the cultivated plants of the South Sea Islands, con-
tain, in addition to numerous well-known facts, many
names of Polynesian plants.

The illustrations of the Caroline and Marian Islands,
and the archipelago of Bonin, are among the most
excellent and richest views in Kittlitz's work ; but the

408 GEOGRAPHICAL BOTANY.

systematic determination of the plants figured is entirely
wanting a deficiency caused by the early death of
Mertens. The vegetation of tropical forests has, in
fact, never been more distinctly represented than in these
landscapes, except by Rugendas. The characteristic
types of the most important physiognomical forms of
tropical foliage are principally found in the following
plates : Bamboo form, indicated by Artocarpm (pi. X) ;
Banana, form expressed by the Rhizophorce of the man-
grove forests (pi. V), and stems of Ficus supported by
aerial roots (pi. VI) ; Cycadese (pi. XI), Palmese (pi. IX,
XVI), MusaceaB (pi. VII), Pandanus (pi. X, XI, XII,
XV), and that of the Tree-ferns (pi. XVI). Of other
physiognomical forms, Lianes (pi. VIII, XV), Freycinetia
(pi. VI), parasitical Ferns (pi. V, VI), Aroidese (pi. VII),
Agave, imitated by stemless species of Pandanus (pi. XI,
XII), herbaceous Ferns (pi. VIII). PI. XIII represents
the savannahs on the Marian Islands ; grass plains with
Casuarina, Cycas, and Pandanus.

Suttor's paper, read before the Linnsean Society, upon
the Forest Trees of New Holland contains, according to
the extracts before us, only well-known facts (Ann. Nat.
Hist., xiii, p. 217). Drummond's Letters from the Swan
River have been continued (Lond. Bot. Journ., 1844,
pp. 263, 300). They contain, for the most part, notices
of individual plants which were transmitted to Hooker.
The extensive herbaria brought by Preiss from the Swan
River have been systematically described in detail in a
separate work, edited by Lehmann, by a number of scien-
tific men, mostly Germans (Plantse Preissianae sive enu-
meratio plantarum, quas in Australasia occidentali et
meridionali occidentali collegit L. Preiss. Ed. Chr.
Lehmann. Vol. i. Hamburgh!, 1844-1845, 8vo). The
coadjutors were Bartling, Bunge, Klotzsch, Lehmann,
Meissner, Miquel, Nees v. Essenbeck, Putterlick, Schauer,
Sonder, Steetz, Steudel, and De Vriese. Summary of
the families treated of, with the enumeration of new
genera, and those containing most species : 247 Legu-

GEOGRAPHICAL BOTANY. 409

minose (Meissn.) ; 63 Acacia, 10 Chorozema, 15 Gom-
jpholobium, 11 Jacksonia, 23 Daviesia, 15 Gastrolobium,
10 Bossi<ea, Rosaceae 1 (N.), Chrysobalaneae 1 (N.) ;
161 Myrtaceae (Sch.), 15 Verticordia, 14 Calycotlirix,
Sym.phomyrtus n. gen., 15 Eucalyptus, 33 Melaleuca,
10 Beaufortia, 15 Calothamnus ; 3 Halorageae (N.) ;
Onagrariae 1 (N.); 2 Oxalideae (Steud.); Lineae 1 (Bartl.);
6 Geraniaceae (N.) ; 2 Zygophyllaceae (Miq.) ; 25 Dios-
meae (Bartl.) ; Boronia 15; 12 Euphorbiaceae (Kl.) ;
Trachycaryon (Croton sp. Lab.) ; Cafyptrostigma (Croton
sp. Lab.); Lopadocalyx n. gen.; 3 Stackhousiaceae (Bg.);
22 Rhamnese (Steud.) ; Pomaderris 10 ; Cryptandra 10 ;
13 Pittosporacese (Putterl.) ; 17 Poly gal aceaa ; Come-
sperma (Steud.) ; 15 Tremandraceas (Steetz) ; 11 Tetra-
theca ; Platytheca n. gen.; 10 Sapindaceae (Miq.);
Olacineae 1 (Miq.) ; Hypericineae 1 (N.) ; 32 Byttne-
riaceae (Steud.) ; Thomasia 19; Fleischeria n. gen. ; 11
Malvaceae (Miq.) ; Phytolaccaceae 1 (Lehm.) ; 5 Caryophyl-
laceae (Bartl.) ; 5 Portulaceae (Miq.) ; Tetragonella n. gen.;

2 Mesembryanthemeae (Lehm.) ; Frankeniaceae 1 (N.) ;
20 Droseraceae (Lehra.) ; Drosera 17 ; 8 Cruciferae (Bg.) ;
Monoploca (Lepidii sp. D. C.) ; 6 Ranunculaceae (Steud.);
44 Dilleniaceae (Steud.); Hibbertia 26; Candollea 11 ;

3 Crassulaceae (N.) ; Cephaloteae 1 (Lehm.) ; 8 Loran-
thaceae (Miq.) ; 31 Umbelliferae (Bg.) ; Platysace n. gen. ;
ScJicenoltena n. gen.; 99 Epacridaceae (Sond.) ; Astro-
lorn a, Brachyloma n. gen. ; Leucopogon 47 ; Andersonia,
14 ; 3 Primulaceae (N.) ; 8 Lentibulariae (Lehm.) ; 6
Scrophulariaceae (Bartl.) ; 5 Solanaceae (N.) ; 5 Convol-
vulaceae (De V.) ; 5 Boraginaceae (Lehm.) ; 8 Myopori-
naceae (Bartl.) ; 2 Verbenaceae (Bartl.) ; Avicennieae 1
(Miq.) ; 25 Labiatae (Bartl.) ; Colobandra 6, n. gen. ;
Anisandra n. gen. ; 6 Gentianaceae (N.) ; Apocynaceae
1 (Lehm.) ; 5 Loganiaceae (N.) ; 4 Rubiaceae (Bartl.) ;
69 Stylidieae (Sond.) ; Stylidium 64 ; Coleostylis (Stylidii
sp. Benth.) Forsteropsis n. gen. ; 18 Lobeliaceae (De V.) ;
Lobelia 17; Vlamingia n. gen.; 59 Goodenovieae (De V.);
Dampiera 15 ; Scaevola 27; 101 Synantheraceae (Steetz) ;

GEOGRAPHICAL BOTANY.

Jurybia 11; Gymnoyyne n. gen.; Silphiosperma n. gen.;
onolepis n. gen. ; Pacliysurus n. gen. ; Chrysodiscus

-^n. gen. ; Chthonacephalus n. gen. ; Anisolepis n. gen. ;

tPterochceta n. gen. ; Siemssenia n. gen. ; Hyalosperma
n. gen.; Schcenia (Heliclirysi sp.); 2 Plantaginacese (N.);
208 Proteacese (Meissn.), PetropJdla 21, Isopogon 15,
Adenantkos 10, Conospermum 17, Grevillea 29, Hakea
46, Banksia 19, Dryandra 22 ; 16 Thymeleaceae ; Pz-
^efe (Meissn.) ; 7 Lauriiiacese, Cassyta (N.) ; Nyeta-
ginese 1 (N.) ; 6 Polygonaceae (Meissn.) ; 14 Amarantaceae
(N.) ; Trichinium 19; 14 Chenopodeacese (N.) : Urti-
cacea3 1 (N.) ; 9 Casuarinese (Miq.) ; 2 Conifers (Miq.) ;
Actinostrobus n. gen. ; Cycadacese 1 ; Macrozamia (Lehm.)
Hence at present about 1450 Dicotyledons.

Gunn has addressed some botanical letters from Van
Diemen's Land to the editor of the ' London Journal of
Botany' (1844, pp. 485-96). He describes an excursion
to the western highlands of the island, and gives state-
ments of the localities of rare plants, with a more detailed
report upon a new species of Eucalyptus (E. Gunnii)
(Hooker, //.), which in December and January contains
a large quantity of a saccharine and fermentable juice,
whence it is called by the colonists the cider-tree. As it
forms extensive mountain-forests, it will probably here-
after become an important product of Tasmania. Harvey
has described some new Alyce from Van Diemen's Land
(id. pp. 407, 428) ; amongst them the Rhodomeleaceous
plant, Pollexfenia, which is also indigenous to the Cape.
Contributions to the Flora of New Zealand : Catalogue
of a Collection of Plants from New Zealand, by Stephen-
son, determined by J. D. Hooker (Lond, Journ. Bot.,
1844, pp. 411-18) it contains but few new species;
Hepaticse Novse Zeelandiae et Tasmaniae, by J. D.
Hooker and Taylor (id. pp. 556-82) ; Diagnoses of New
Zealand Plants, by Raoul, preliminary to his illustrated
work, which was published in 1846 (Ann. Sc. Nat.,
1844, ii, pp. 113-23), with the new genera: Ileodictyon
(Fungi), Pukafaria (Corneacese ?), and Tctrapalhea
(Passifloreae) .

GEOGRAPHICAL BOTANY.

Colenso's Botanical Diary of his travels during sever r
months through the little known interior of the norther,,
islands of New Zealand (Lond. Journ. Bot. 1844, pp. 4
62) contains numerous localities of, and reports upon,\
newly-discovered plants which have not yet been made\
public, but will be described in Dr. Hooker's illustrated
work.

The first three parts of the latter work have appeared ;
they contain a general introduction upon the botanical
characters of high latitudes of the southern hemisphere,
and also the commencement of a flora of the Auckland
Archipelago (The Botany of the Antarctic Voyage of
H. M. Discovery Ships Erebus and Terror, under the
command of Sir J. Ross ; by Jos. Dalt. Hooker. Parts
i-iii, London, 1844, 4to.) Being, during the summer,
almost always either in high barren latitudes, or on the
open sea, Hooker had but little opportunity of collecting
other than such plants of the antarctic flora as flowered
in the winter or spring. But he considers this defect,
which concerns the copiousness of the materials which he
collected, as of little consequence, as he was in the favo-
rable position of being able to make use of the botanical
results obtained in all the earlier British voyages to the
South Pole, but of still less, in consequence of a climatal
peculiarity which he developes in the introduction, and
regards as the most characteristic feature of the antarctic
vegetation. He was surprised on finding at Kerguelen's
Land, the same plants in flower which Cook had met
with at other seasons, and this result he subsequently
found to occur generally. The vast preponderance of
water in the high southern latitudes produces an uni-
formity in the distribution of heat throughout the year,
and the more we approach the pole the more distinctly
does this appear to increase. The seasons there are not
distinguished, as in the north, by their temperature, but by
scarcely anything more than the variation in the amount
of light ; all the months are cold, but the thermometer
varies, as in the tropics, within narrow limits. In the

412 GEOGRAPHICAL BOTANY.

region of floating icebergs, between 5 5 and 65 S. lat.>
seldom a day occurred during the summer in which the
temperature rose or sunk beyond the limits of c. and
6'6 c. South winds, with much snow, alternate
there with aerial currents from the north, which being
loaded with aqueous vapour, incessantly diffuse white
fogs of indescribable density over the surface of the ocean.
These precipitations are also formed on islands situated
in the vicinity of this region, throughout the year, by the
admixture of the winds from the land and sea depriving
them of their solar climate, and for the most part pre-
venting the change of temperature dependent upon the
position of the sun. A climate so inhospitable and
uniform excludes any variety in the forms of plants, but
confers a luxuriance of growth upon the indigenous
plants, of which the arctic regions must necessarily be
deprived, because their vegetation is subjected to a pro-
longed winter-sleep. This is so remarkably the case,
that notwithstanding the differences in the climatic con-
ditions, most of the genera and forms of the antarctic
agree with those of the arctic flora in the most important
points, excluding only the Auckland Islands, which ap-
pear to belong to the same primitiveformation with New
Zealand. But notwithstanding this similarity of the
types, the species of the southern district are peculiar ;
which could not have been expected to be otherwise in
the case of islands, not only separated climatally to such
an extent, but are also situated beyond the reach of all
continents, the oceanic currents of which usually plant
the waste shores. Many antarctic species indicate their
endemic origin by the limited district through which they
are distributed in the region itself. However, the special
botanical results of Hooker's voyage, the description of
which far excels his former communications in fulness
and arrangement of the matter, are reserved for the next
Annual Report. The Cryptogamia have also been partly
described in the ' London Journal of Botany' for 1 844,
including 72 Hepaticae from the Auckland Islands, by

GEOGRAPHICAL BOTANY. 413

Hooker and Taylor, (p. 366 ;) 66 sp. more from the
Falklarids, Cape Horn, and Kerguelen's Land, by the
same author, (p. 454 ;) 73 antarctic Jungermanniee, by
Hooker and Wilson ; with the new genera Lophiodon and
Hymenodon (p. 533), and 151 antarctic Lichens, by
Hooker and Taylor, (p. 632.)

Dr. Hooker paid particular attention to the distribu-
tion of the Alyce floating in the high latitudes of the
Southern Ocean. (Antarct. Voy., Introduct.) Macro-
cystis and Urvillea were found common as far as the
northern limit to the floating ice, in one instance they
extended to 64 S. lat. ; but they usually disappeared
much sooner, e.g. south-east of America, below 55S.
lat. But in the latter meridian a new form of Alga
appeared below 63 S. lat., which although previously
found in D'Urville's expedition, has since been described
as Scytothalia Jacquinotii. Here, near the coast of
Palmer's Land, on Cockburn's Island, (64 S. lat.), no
Phanerogamous plants were met with, only 20 Crypto-
gamia. These appear to be the last forms of plants in
the direction of the antarctic pole ; for even the Algae are
absent on that continental coast upon which the active
volcano Erebus and .the extinct volcano Terror are
situated, and where the soil at the level of the sea ap-
peared for the first time entirely deprived of vegetation,
a sight never before witnessed, and from which nature
appears to have preserved even the highest latitudes of
the north.

REPORT

ON THE PROGRESS OF

GEOGRAPHICAL AND SYSTEMATIC
BOTANY,

DURING THE YEAS, 1845.

BY DR. A. GRISEBACH,

EXTRAORDINARY PROFESSOR IN THE UNIVERSITY OP GOTTINGEN.

GEOGRAPHICAL AND SYSTEMATIC

BOTANY.

THE consideration that the greater number of literary
productions in the province of Systematic Botany are
connected with the working out of individual local floras,
and ought therefore to have been noticed in the preceding
botanico-geographical Annual Reports, has convinced the
author of them, that by altering the arrangement of the
matter, with an appropriate limitation of the descriptive
notices, Systematic botany also may be made to enter into
these notices, without the allotted space being exceeded.
The year 1845 has moreover proved comparatively poor in
botanico-geographical results, so that the present period
appears appropriate for a first attempt at enlarging the
botanical Annual Reports in correspondence with this view.
Hence by comprising, in combination with those upon
Vegetable Physiology, the entire domain of Botany, they
will now obtain a completeness corresponding to that of
the Zoological Reports, and, as I hope, will acquire more
practical utility. An important limitation will, however,
still exist in the Botanical Reports, viz. that both the
descriptions of the plants, as also the notices of individual
species of genera already known, are excluded from the
sketch given of the systematic works ; not only, however,
does want of space compel us to omit these considera-
tions, but it would also be superfluous to repeat here
what is annually done in so admirable a manner in several
botanical periodicals and repertories.

418 BOTANICAL GEOGRAPHY.

,4. BOTANICAL GEOGRAPHY.

R. B. Hinds has continued his general observations
upon botanical geography (see Annual Report for 1842)
during the past year (Memoirs on Geographic Botany,
Ann. Nat. Hist. vol. xv) ; like the former, however,
they contain little more than already known facts and
views, not unfrequently mingled with errors, both as
regards the facts and deductions.

The present papers contain, e. g. estimates of the whole number of plants
known ;* remarks upon centres of creation, the existence of which the
author denies ; on the distribution of certain families ; on the mean area of
the extension of each species ; elements for the comparison of two floras ;
on physiognomy, &c. I shall only enter upon the consideration of one of
these views, and that because it places a simultaneous work by Forbes, re-
markable for its originality, in a proper light. The old hypothesis of a single
centre of creation, from which all plants upon the earth were distributed, as
also the later supposition, that tliis migration of organisms originated in one
or a few centres, Hinds opposes by the general law, that wherever plants
met with the conditions necessary for their existence, the present vegetation
was originally produced. He does not admit, in opposing every migration of
plants, even such variations in the original state as would cause the extirpation
of individual species, and their disappearance from among the number of
living organisms ; whilst such an event, e. g. in the case of the endemic
plants of St. Helena, has been as satisfactorily determined as in that of
Didus ineptns. The historical change in the constituents of forests, and the
migration of individual plants which is in progress before our eyes, and is
not merely produced by man, are incompatible with a law expressed so
generally. The fact that certain islands in the Indian Ocean, as e. g. Darwin
showed, contain only plants which have been washed on shore, by wliich they
are thickly covered, whilst the islands in their vicinity possess an endemic
vegetation, contradicts the supposition of the existence of a generally diffused
productive force, or at least limits it to distinct creative epochs. When we

* Hinds estimates the number of known plants at 89,170; those existing
on the earth at 134,000 species. His statements are founded upon the
number of species contained in the first four volumes of De Candolle's
' Prodromus.' These amount to 20,100 ; of which 3875 are Leguminosce,
1631 Rubiaceje, 1009 Umbellifera, 990 Cruciferje, 759 Caryophyllaeeaj, 715
Myrtacese, &c.

BOTANICAL GEOGRAPHY. 419

reflect on the well-known facts which Hiuds brings forward, without either
certainty or accuracy of detail, as the foundation of his opinions, they leave
room for the formation of other hypotheses besides his. His laws are as
follows : 1. In proportion as the districts of vegetation are further separated
from each other by the sea, so is the number of species of plants common to
them less. Hence the large number of species common to the three large
portions of the earth in the arctic zone, and hence the greater the contrast,
in comparing the floras of corresponding climates, the further south we pro-
ceed, the various portions of the earth in the southern hemisphere being
further separated from each other. 2. If we divide the whole earth into
six floral districts which would certainly be arbitrary enough we obtain
for each almost exclusively endemic species, and we may add, that the same
result would hold good were we to admit more than thirty districts. 3. In
different natural floras under corresponding climates, we certainly find
similar forms, but not the same species. 4. In some islands the vegetation
is entirely endemic ; these cannot, therefore, have obtained their plants by
migration from external sources, &c. All these and similar facts certainly
are opposed to the migration of plants from a single point of the earth's sur-
face to all the rest, and which scarcely any naturalist now believes ; but
there is a broad interval in the argument, which has not yet been filled up
with facts, between these considerations and the assertion, that centres of
creation do not exist, but that every point has produced the plants it pos-
sesses. We know that some regions of the earth contain many more endemic
species than others, without the soil or climate being sufficient to explain
this increase. The number of endemic forms diminish in the direction of
any climatal boundary, as it were, like the radii of a circle, in the centre of
which a centre of creation is situated ; hence, e. g. in Europe, we may speak
of western, eastern, or southern forms of plants, which gradually disappear,
one after the other, towards the east, the west, or the north. There does
not appear to be any other difference between an island which possessed
endemic plants only, as St. Helena, and a continental district, which, like
Spain or Illyria, abounds in endemic plants, than that other plants which
have migrated from external sources are associated with the latter, which
could not readily occur in the former case, on account of the distance of
any continent. On reviewing all the known facts, and seeking for the
simplest theory by which their relation may be explained, we are com-
pelled to adopt the supposition of the existence of as many centres of creation
as there are districts of endemic plants upon the globe. The difficulty of
determining in each case the original centre of creation, on account of the
mixture of the groups in the wide and connected districts of continents, is
so great, that it must always remain the main object of botanical geography.
It is only the problem of the groups of creation that gives this science a
peculiar importance, and raises it above the imputation of being an aggre-

420 BOTANICAL GEOGRAPHY.

gate of heterogeneous laws belonging to different sciences ; for in this light
only does it present a definite and independent method of investigation, a
progressive course of development.. Commencing with observations upon
the geographical area of each individual species of plant, the object of
botanical geography is first to determine what limits to this distribution have
been placed by the composition of the soil or the subdivision of the continent.
It then points out the climatal sphere of the species, and if it ascertains,
after this twofold limitation, that the natural area is more contracted than is
accounted for, the geological problem presents itself what has not been the
result of soil and climate must depend upon historical causes, the history of
the earth. If the same soil and the same climate have produced only similar
but not identical forms, this refers us to a creative act of a different kind,
therefore to a geological epoch.

In connexion with this combination of geological and botanico-geographical
investigation, Forbes has made an attempt of a different kind, viz. the appli-
cation of the distribution of plants to geological deductions (Report of the
Meeting of the British Association, held at Cambridge, Ann. Nat. History,
xvi, p. 126). On comparing the specific centres of the endemic plants of
Great Britain, i. e. the centres of their geographical areas, it is evident that
the flora of the greater part of the surface of the country belongs to that of
Germany. The specific centres of the few species peculiar to the British
Isles occur in the same region. In addition to this principal area, four
smaller districts of vegetation may be distinguished according to the same
law : 1. The mountainous districts of the west of Ireland contain a number
of plants in common with the north-west of Spain and the Pyrenees. 2. The
south country, Devonshire, Cornwall, and the Channel Islands, in common
with the west of Trance. 3. The south-east of England, especially its
chalky districts, with the north of France. 4. The mountains of Wales, the
north of England and Scotland, with the plains of Norway. Forbes does
not .consider this connexion as explicable by soil and climate, and therefore
seeks for geological causes, in conformity with the above law. He believes
that these are to be found in a former connexion of Great Britain with the
continent, probably existing in earlier geological periods, especially during
the tertiary period : not that this connexion, made use of for his explanation,
has been geologically determined ; but he endeavours to support his geological
hypotheses by these botanico-geographical relations. Following up this
design, which is certainly not free from objection, Forbes then not only
maintains generally these connexions of land, but by supposing former eleva-
tions and subsidence of the soil, arrives at certain views regarding the series
of changes which have occurred ; in fact, he distinguishes the floras according
to the periods at which they were formed. I should have no hesitation my-
self in granting, that when two different floras really belong to the same soil
and climate, by far the simplest hypothesis is to attribute their origin to

BOTANICAL GEOGRAPHY. 4.21

different geological epochs ; but if, as I believe to be the case, climatal con-
ditions sufficient to account for the above distribution of British plants were
present, the error would not lie in the method, but its application, which has
led Forbes to the following results. According to his view, the districts of
vegetation distinguished above correspond to as many geological eras, so that
the flora of the west of Ireland would be the oldest, that of the mountains
the fourth, and that related to Germany the youngest. The first mentioned
descend from a time when a chain of mountains, running across the Atlantic
Ocean, connected Ireland with Spain ; this would explain its difference from
the vegetation of the mountains, although it would still correspond with the
mountain character. Moreover, in the second and third periods the English
Channel was closed, first towards the west, then towards the east also, by the
connexion of land, and thus the distribution of French plants in England was
occasioned. Forbes explains the alpine flora of the mountains by means of
Agassiz's glacial period ; the mountain summits of Britain were then low
islands, extending to Norway, and were clothed with an arctic vegetation,
which, after the gradual upheaval and consequent change of climate, became
limited to the summits of the newly-formed and still existing mountains.
Lastly, the bed of the North Sea itself was upheaved, and extensive plains
laid dry between England and Germany, upon which the elk and other extinct
quadrupeds lived, and over which the plants of Germany migrated ; until at
last the sea, in consequence of fresh depression, flowed back, after the im-
portant object of transplanting Roses and RuU beyond the ocean had been
accomplished. Further than this, hypothetical views could not easily be
carried, and I have translated them almost entirely here, only because Forbes
appears desirous by this paper of opening a new path in botanical geography,
for this first lecture has since been followed by others. The criticism of his
undertaking lies simply in the denial of one of the first statements with
which he commences : actual natural forces, the sea, rivers, currents of air,
by which seeds are diffused, or animals, and even man, are, in the majority
of cases, insufficient means for effecting the migration of plants across the
British seas. I maintain that these forces are quite sufficient, provided the
imported seeds meet with a suitable climate and proper soil. Those western-
Europe plants, which, being produced through the agency of the coast -
climate of the Atlantic, and, according to the degree of this dependence,
becoming distributed sometimes to a greater, sometimes a less distance
within the continent, the author refers in the latter case to Spain, in the
former to France. are not met with equally on the coast-line of the con-
tinent, but are often absent from wide tracts, the soil of which is not
favorable to their growth ; when e. g, we do not find Erica cinerea anywhere
from the Rhine to the Fjord of Bergen, who would, in this case, suppose
that former connexions of land had disappeared, when for the most part the
connexion still exists, without contributing to the distribution of this shrub ?

422 BOTANICAL GEOGRAPHY.

Since the Alps contain so many alpine species in common with the arctic
regions, it is still more easily seen how little the continent situated between
these two terminal points serves to explain these agreements. The plains
which, without this alpine attire, extend e. g. from Cola to the Carpathians,
are, however, less adapted to the transport of foreign plants than a sea which
rapidly carries over the seeds in its current. Or when Forbes has recourse
to the glacial period in explaining the diffusion of the arctic plants : how
will he account for so many central European species of Sierra Nevada or
Pindus traversing the extensive tracts of land which separate them from
their centre of creation ? How, by the most complex dislocations, will he
bring the Minuartice and Queritf into geological connexion, which do not
nourish anywhere between Castile and the Crimea? There is no reason
why water should form a greater obstacle to the distribution of plants than a
soil which does not conduct them ; extensive oceans certainly form barriers,
when there is no current to carry them across, or when the climates of the
two coasts are dissimilar.

A. Erman has written a paper on the periods of vege-
tation in different climates (Arch, fur Russian d, Bd. 5,
pp. 617-40).

He examines the question, of what relation the stages of development
of vegetation hold to the temperature, at which in different latitudes they
appear in the same species. His investigations lead to the negative result,
that a law communicated to him conjecturally by Quetelet is unfounded.
It consisted in the assumption that similar stages of development occur in
two different places, when the sum of the squares of the diurnal temperature
from the commencement of the period of vegetation is the same for both. He
shows, also, that the stages of development and the sum of the temperature
acting upon them in different places, are by no means in direct proportion.

We must mention a remark made by J. D. Hooker in
regard to botanico-geographical physiognomy (On Fitchia,
in Lond. Journ. of Bot., 1845, p. 640).

On many remote islands possessing endemic floras, we find woody plants
belonging to the family of Synantheracese, which contribute essentially to
the character of the districts, and belong to peculiar genera, of which re-
presentatives do not occur on the continents. The following sketch will
serve to illustrate this :

St. Helena contains 4 gen. 10 sp. of Synantheracea?, all woody plants ;
Juan Fernandez 8 17 of of which 3 gen. and

12 sp. are woody plants ;
Gallapagos contains 13 gen. 21 sp. of Synantheraceae, of which 3 gen. and

8 sp. are woody plants ;

New Zealand contains 30 gen. 60 sp. of Synautheracea3, of which 8 gen.
and 14 sp. are woody plants.

BOTANICAL GEOGRAPHY. 423

Elizabeth Island, which belongs to the botanical district of the South
Sea Islands, in the southern hemisphere, but is situated nearer to the island
of Juan Fernandez and the continent of America than the others, also con-
tains the new arborescent Cichoraceous genus, Fitchia, whilst the other
islands of this archipelago do not contain similar forms of plants.

I. EUROPE.

Of Ledebour's Flora Rossica (see the Annual Report
for 1841 and 1843), the 6th part appeared in 1845,
and the 7th in 1846 (vol. ii, part 2).

The statistical proportions of the families treated of in them are as fol-
lows : Synantheracese, 890 sp. The Vernoniacese are only represented by
the Caucasian Gundelia ; of the Eupatoriae, in addition to the West-Euro-
pean genera, Nardosmia and 7 arctic species ; the Asteroidese comprise the
exclusively Asiatic genera Turczaninoma, Calimeris, Arctogeron, Diplopappus,
Rhinactina, Myriactis, Brachyactis, Dicrocephala, Karelinia, Eclipta, and
Siegesbeckia, which extends as far as the Crimea ; among the Senecionidae,
including the Heleniee, RicMeria and Cancrinia, Brachanthemum, one of the
Chrysanthemeae from Siberia, Valdheimia from Altai, Cladochaeta and
Amblyocarpum from the Caucasus, and Senecillis from Podopolien. The
genera most abundant in species are Artemisia (83 sp.), Senecio (52 sp.),
Achillea (31 sp.), Pyrethrum (29 sp.) ; among the Cynaracese, including
Acanthocephalus and Haplotaxis (3 sp.), and Ancathia from Altai, Alfredia
(4 sp.) from Siberia, and Cousinia (20 sp.) and Acroptilon from the Steppes,
and Acantholepis, Chordinia, and Oligochceta from Armenia. Those con-
taining most species are Centaurea (61 sp.), Cirsium (51 sp.), Serratula with
Jurinea (36 sp.), Samsuria (32 sp.) ; the Cichoraceae contain Heteracia and
Microrhynchm from the Steppes, Asterothrix from the Caucasus, Intybellia
from the Crimea, Youngia (5 sp.) from Armenia and Siberia, Ixeris and
Nabalus from Siberia, Apargidium from Sitcha ; and of larger genera, Hiera-
cium (25 sp.), Crepis (23 sp.), Scorzonera (19 sp.), Lactuca (17 sp.), Trago-
pogon (17 sp.) ; Lobeliaceas, 2 sp. ; L. dortmanna, and in Eastern Siberia
L. sessilifolia; Campanulaceae, 66 sp., containing Michauxia and Symphyandra
from the Caucasus, Platycodon from Darien ; the genera containing most
species being Campanula (36 sp.) and Adenophora (10 sp.) ; Vaccinias,
11 sp., including 4 sp. from Sitcha, 1 sp. from the Aleutian Islands, and
V. arctostaphylos from the Caucasus ; Ericese, 36 sp. ; 2 Rhododendra and
Azalea Pontica, confined to the Caucasus, extending hence to Dombrowitza
in Lithuania, 4 sp. of Cassiope, Eryanthus, 2 sp. of Amothamnus, 5 sp. of
Rhododendron confined to Siberia, 2 sp. of Cassiope, Menziesia, 1 sp. of

424 BOTANICAL GEOGRAPHY.

Phyllodoce, Kalmia, and Cladothamnus confined to Sitcha ; Pyroleae, 7 sp.,
corresponding to the German species ; Monotropeae, 1 sp.

The 5th and 6th parts of Trautvetter's illustrated work
1 Plantarum imagines Moram Rossicam illustrantes/ Mo-
nachii, 1845, 4to, (see the preceding Annual Report)
have appeared, containing pi. XXI XXX.

The Academy of St. Petersburg has commenced the
publication of Contributions to the Botany of the
Empire of Russia' (Part 1, Petersburg, 1844, 30 pages
8vo. ; part 2, 67 pages and 6 plates ; part 3, 56 pages ;
part 4, 93 pages; ib. 1845).

The first part contains a local flora of the province of Tambow (incomplete,
containing 312 sp.), forming Ruprecht's fourth contribution to the flora of
St. Petersburg. The same author has published an account of the Eerns
and Charge of the empire of Russia in the third part ; it also describes some
new Terns from Siberia, Mongolia, and American Russia, as also of the
Charse from the Soongari.

The second part, in which Ruprecht has described his botanical travels in
the extreme north of European Russia, is of more general interest. In the
unfavorable summer of the year 1841, he collected in the eastern part of the
province of Archangel, especially Mezeu, the peninsula of Kauin, and the
island of Kalujew. The natural character of the country differs from that
of Scandinavian Lapland in the circumstance that the forest-limit recedes
almost to the vicinity of the arctic circle ; hence extensive, low, treeless
plains are spread along the arctic ocean. Thus the pine forests are en-
tirely absent at Kanin (excepting a wood composed of Abies, situated below
67} N. lat., and which is now dying) ; they cease at the Indega river, about
fifteen English miles from the sea, and scarcely extend over the arctic circle
beyond Petschora. The cultivation of barley and potatoes also is only
carried on as far as the city of Mezen. The forests are succeeded north-
wards, first by a zone of low birches and willow-shrubs, the dwarf birch
with the arctic Ericacea? follow next, and lastly, with the latter the continuous
turf of the alpine regions terminates. A few Ranunculaceae, Saxifrages,
and Grasses, which only partially cover the soil, are all that are subsequently
met with. In these travels 342 Phanerogamic plants altogether were col-
lected. They also differ from those of the flora of Scandinavian Lapland,
in containing a large proportion of species which are riot Scandinavian.
Eleven new species, which are illustrated by figures, belong to the genera
Ranunculus, Viola, Parnassia, Salix, and Poa (7 sp., 1 sp. of each of the
others).

Czerniaiew has published some scattered remarks on

BOTANICAL GEOGRAPHY. 425

the influence of climate upon the vegetation of Ukraine,
at the same time introducing the description of some new
Fungi (Bulletin des Naturalistes de Moscou, torn, xviii,
part 2, pp. 132-57).

Many plants are excluded by the low Isochimena, whilst the high summer
temperature appears favorable to the culture of Maize and several Cucurbi-
tacese, by which also the author endeavours to explain the remarkable fact,
that the berries of Solatium nigrwm in Ukraine lose their narcotic principle,
and when ripe become saccharine and eatable. The forests and fields there
are protected from the persistent aridity of the summer, which acts to so
great a degree upon the adjacent steppes, by the soil of humus, which is
from 10 to 15 feet in depth (Tscherno Sem; compare Ann. Rep. for 1843.)
Hence the principal forest trees which thrive there are such as send out
deep roots, as the oak^the lime, elms, and pear trees ; the red fir (P. Abies),
which predominates on the shallow soil of Scandinavia, is unknown in
Ukraine, and the ash is frequently killed during the dry season. The deep
soil of humus causes several herbaceous plants to grow there to an unusual
height ; Cephalaria Tartarica is found 9 feet, Delphinium elatum 5 6 feet
in height ; Thistles and Umbelliferse are usually twice the size of those of
other regions ; of the Fungi, the pileus of Polyporus and Leuzites is found
three feet in breadth ; and the new Morchella alba a foot in height. But
the most remarkable object presented by this luxuriant development is the
new Bovista, Lycoperdon horrendum, a spherical Fungus, 3 feet in diameter.
This Fungus, says the author, might really produce no slight amount of
terror ; when in the dark forest it suddenly comes into sight, it makes one
imagine a phantom in white or brown garments and in a stooping attitude.
This black earth of the south of Russia, which produces this luxuriant
growth, must indeed contain a large store of nutritive matter for the vege-
table world ; for barley grows there as in the best districts of England or
Germany, without ever requiring manure. As regards the Fungi of
Ukraine, Czemiaiew attributes the very numerous varieties of their forms
to the paucity of the species of Mosses, Lichens, and Ferns. According to
his observations, Ukraine alone contains more than 1000 Hymenomycetae,
whilst the abundance of Gasteromycetes is still more characteristic. Wein-
mann, in his ' Prodromus/ which was published in 1 836, enumerates 300
Gasteromycetes as existing in the whole of Russia, whilst Czerniaiew has
found almost twice this number of species in Ukraine alone : among them
there are several new forms, and a few new genera.

Weiumann has studied the Mosses of Russia (Bulletin Moscou, torn, xviii,
pt. i, pp. 409-89, and pt. ii, pp. 417-503) ; his new species belong to
Funaria (1 sp.) and Hypnum (4 sp.) Kaleniczenko describes ten new
plants from the south of Russia and the Caucasus (Ibid. pt. i, pp. 229-40) ;

426 BOTANICAL GEOGRAPHY.

2 Umbelliferee (Pimpinella and Pastinaca), 2 Leguminosae (Arthrolobium),
6 Synantheraceae (Inula, 2 sp., Centaurea, 3 sp., and Jurined),

The travels of A. Bravais and Gh. Martins (Bibliotheque
univers. de Geneve, 1845, vol. ii, pp. 147-73) traverse
the north of Scandinavia almost exactly in the same way
as that described by V. Buch in his celebrated work on
the extreme regions of the north, when he returned from
Alten-Fjord, in Finmark, to Tournea, on the Gulf of
Bothnia. But the French travellers believe that they
have measured the limits of vegetation under more favor-
able circumstances ; hence their results must find a place
here. They completed the difficult journey from the 6th
to the 26th of September, 1839 : remarking at the same
time, that partly on account of the water which they had
to pass over, partly on account of the swarms of gnats
met with in Lapland during the summer, which have to
be avoided, September is the only month fit for travelling.

In the forests of Alten (70 N. lat.), the pines were 60' in height, the
birch on an average 45'. On the third day they passed over the upper
terrace of the plateau of Kjolen. Under the name of Nuppivara, it ascends
here to an elevation of only 600 metres ; but it is of the same conformation
as the far more elevated, undulating plateaux of the Langfjelde, which
abound in lakes. The bare soil of the rocks contains only a scanty under-
wood of Betula nana with Empetrum, and Andromeda tetragona, or Salix
Lapponum with Juniperus communis. On the south side, birch forests are again
next met with, but they do not extend above Kautokeino further than to
Karesuando (68 36') ; for from this point the whole country, as far as the
Gulf of Bothnia, is covered with a single continuous pine forest.

Limits of vegetation measured :

Northern slope of Kjolen, in the valley of Alten.

Pinus sylvestris, dense forest 249 metres

isolated dwarf . . . . . 500

Betula pubescens, dense forest 380

in the form of a stunted tree . . 432

local . 534

Southern slope of the Nuppivara.

Belula pubescens 477 m., 480 m.

(The figures in apposition denote the results of different measurements.)
Sorbus Aucuparia 4? 7 m.

BOTANICAL GEOGRAPHY. 427

Dividing line between the rivers of the North Sea and of the Baltic :

Region extending from Kalanito to Suvajervi.
Pinus sylvestris ....... 341 m., 374 m.

Setitla pubescens .... 493 m., 498 m., 520 m., 530 m.

Sorbus Aucuparia . . . . . . . .474m.

At Karesuando.
Pinus sylvestris ........ 410 m.

A list of the Phanerogamous plants occurring at Karesuando, by Leesta-
dius, is inserted in the report of these travels.

A report by Blytt on his botanical journey through the
valley of Valders in Norway, contains, for the most part,
merely copious lists of localities (Bot. Notiser, 1845,
Nr. 1-3). The author, however, in his account of the
calcareous vegetation of Torpe, adds some remarks upon
the influence of lime on the distribution of Norwegian
plants.

Very few peculiarly calcareous plants are found there, and several species
which in other countries are confined to a calcareous soil, grow upon the gneiss-
formations of Norway. Blytt only enumerates the following Phanerogamia as
belonging to the chalk in Norway : Anemone ranunculoides, Trifolium monta-
mtrn*, Libanotis*, Monotropa, Stachys arvensis, Carduus acanthoides*, Ophrys
my odes*, Neoltia nidus avis, and Malaxis Loeselii ; those species only marked
with an * are, according to my knowledge, peculiar to calcareous soils in
other regions, nor are the Lichens and Mosses enumerated, so in every case.
Blytt then criticises Unger's well-known catalogue of lime-plants, and in so
doing, he separates the following species, which grow in Norway on the
gneiss-formation, and part of them on it only : Hepatica triloba, Corydalu
fabacea, Astragalus glycyphyttus, Dryas, Rubus saxatilis, Sorbus aria, Coto-
neaster vulgaris, Saxifraga oppositifolia, Asperula odorata, Pyrola rotundi-
folia, Arctostaphylos alpina, Fagus, Taxus, Convallaria majalis, verticillata,
Polygonatum, Calamogrostis sylvatica, and Brachypodium gracile. Grimmia
apocarpa.. Hypnum Halleri, Lecidea vesicularis and Candida, and Gyalecta
cupularis.

Blytt points out similar differences between Norway and Tyrol, in regard
to those plants which, according to Unger, are more common on a calcareous
soil on the Alps than upon other substrata.

W. P. Schimper has given a description of the Dovre-
fjeld, especially the Mosses found on it, several new species
even of which he has discovered on this soil, which has
been so frequently described (Ratisbon Flora, 1845,
pp. 113-28.)

428 BOTANICAL GEOGRAPHY.

Swedish works upon the botanical topography of Scan-
dinavia : Anderson, Plantae vasculares circa Quickjock
Lapponiae lulensis (Upsal, 1845, 8vo, 36 pages) ; contains
356 sp.; Lagerheim and Sjogren, Botanical Remarks made
during a journey from Stockholm to Snaasahog in Jemt-
land, in 1844 (Bot. Notiser, 1845, Nr. 11) ; Schagerstrom,
Conspectus vegetationis Uplandicae (TIpsal, 1845, 8vo,
83 pages) : contains 870 sp. ; Lindeberg, an Excursion to
the Malar Lake (Bot. Notiser, 1845, Nr. 12) ; Lindgren,
Notices upon the Vegetation of the Wener Lake (ibid.),
with a description of some new pileate Fungi ; Lindeberg,
on the Country around Grenna on the Wetter Lake (ibid.
Nr. 4). Systematic contributions to the Mora of Sweden :
Anderson, Salices Lapponise, cum figuris 28 specierum
(Upsal, 1845, 8vo, 90 pages) : described according to
Fries's views ; Lund, Conspectus Hymenomycetum circa
Holmiam crescentium (Christiania, 1845, 8vo, 118 pages).

On the botanical topography of Denmark : Petit, Re-
marks on the Vegetation of the south-west of Zealand
(Kroyer's naturhistor. Tidskr., second series, vol. i) ;
J. Lange, on the Vegetation of Laaland and Falster (ibid.)
In the case of a tolerably large number of the plants
mentioned here, the north limit to their distribution is
situated on these islands.

Watson is preparing some new works on the botanical
geography of Great Britain, and has made a report upon
the plan adopted in them (Lond. Journ. of Bot. 1845,
pp. 199-208). He very properly takes care to separate
the topographical details of the localities from the more
general investigations, which are of real scientific interest.
The two botanical regions which he distinguishes in Great
Britain, he denominates the Agrarian and the Arctic ; the
area of the region of the Cerealia coincides with the dis-
tribution of Pteris aquilma.

Further contributions to the botanical topography of
Great Britain : Balfour, on some Excursions on the Scottish
Peninsula of Cantire, and the Island of Isla, one of the
Hebrides (Ann. Nat. Hist, xv, pp. 425-6) ; Gardiner, on

BOTANICAL GEOGRAPHY. 429

the Highlands of Braemar (Botanical Rambles in Braemar,
Dundee, 1844), described in a picturesque style; Moore,
on the rare Plants of Yorkshire (Report of the British As-
sociation held at York, pp. 70-1) ; Andrews,on the Island
of Arran and the West Coast of Ireland (Lond. Journ.
of Bot. 1845, pp. 569-70).

Local British Floras : On the county of Cork in Ireland
(The Botanist's Guide for the County of Cork : Contri-
butions towards a Fauna and Flora of the county of Cork,
London, 1845, 8vo), contains 885 Phanerogamia, and 936
Cryptogamia ; Jenner, on the country around Tunbridge
Wells, in Kent (A Flora of Tunbridge Wells ; Tunbr.,
1845, 8vo), includes also the Cryptogamia.

Systematic works on British plants : Bell Salter, three
new species of Eubus (Ann. Nat. Hist, xv, p. 305) ; Bab-
ington on Cuscuta (ibid, xvi, pp. 1-3), containing figures
of C. trifolii and C. approximata Bab., the latter brought
over from the East Indies, with seeds of Melilotus; Parnell,
on the Grasses (Descriptions of the Grasses of Great
Britain, illustrated by 210 figures) ; Spruce, on some
newly-discovered Mosses (Lond. Journ. of Bot., 1845,
pp. 169-95); 23 Jungermanniae with 4 new species;
Taylor, on 6 Jungermanniae new to great Britain, (ibid,
pp. 276-8,) amongst them one new species; Salwey, the
rare Lichens of Wales (Ann. Nat. Hist., xvi, pp. 90-9) ;
Hassall, a History of the British Fresh-water Algae, in-
cluding the Desmideae and Diatomaceae, with upwards of
100 plates (London, 1845, 2 vols. 8vo). The Phytologist
is continued. The following collections of dried plants
must be mentioned : Salicetum Britannicum auct. Leefe
(see Ann. Rep. for 1843), fasc. 2, see the critical remarks
by Sonder (in Ann. Nat. Hist, xv, p. 275); M'Calla,
Algae Hibernicae, vol. i (Dublin, 1845, 4to), with 50
species; Ayres, Mycologia Britannica (London, 1844),
containing 50 sp., may be regarded as a continuation of
Berkeley's work.

Van den Bosch has published the third part of his
Flora of Zealand (see Ann. Rep. for 1842), containing

430 BOTANICAL GEOGRAPHY.

the Lichens arid some additions (V. d. Hoeven, Tyd-
schrift, vol. xii, pp. 1-22), e. g. in the coast districts of the
Netherlands : Cerastium tetrandrum, Trifolium subterra-
neum, and Centaur ea nigra, are found common, also Salix
holosericea, Car ex trinervis (C. rigida Fl. Leyderis), and
Zygodon viridissimus. The contributions to the Crypto-
garnic Flora of the Netherlands, by Dozy and Molkenboer,
have been continued (ibid. pp. 257-88) : on the Fungi,
containing among them some new species which are
illustrated by figures. General works on the Flora of
Germany : Reichenbach's Icones, vol. vii, Dec. 5-1 0, with
the Naiadeae, Alismacese, Hydrocharidaceae, Nymphaeaceae,
and a supplement to the Grasses ; Sturm's Flora, sect. 1,
parts 89, 90, principally containing species of Viola and
Labiatae ; V. Schlechtendal and Schenk's illustrated work,
vol. vi; Lincke's Publication, parts 50-9; Koch's synopsis,
ed. 2, fasc. 3 (Lips. 1845), containing the Ferns, with
appendices and the Index : an abridgment of this work
appeared as a pirated edition under the false name of
Herold ; Nees v. Esenbeck's Genera Plantarum Florae
Germanicae, continued by Putterlick and Endlicher, fasc.
24 (Bonn, 1845, 8vo). Special systematic works on the
Flora of Germany : Sauter's new Contributions to the
Flora of Germany (Ratisbon Flora, 1845, pp. 129-32) :
unimportant notices, with the diagnosis of a new Riccia ;
Perktold, the Hypna of Tyrol (Neue Zeitschr. des Ferdi-
nandeums, Bd. 11) ; Rabenhorst's Cryptogamic Flora of
Germany (see the preceding Ann. Rep.) vol. ii. pt. 1,
containing the Lichens ; Roemer, the Algae of Germany
(Hanover, 1845, 4 to; with 11 plates), confined to the fresh-
water Algae, principally the forms found by the author
on the upper Hartz, and imperfectly illustrated by bad
lithographic drawings ; arranged according to Kiitzing's
system, but paying no attention to the history of develop-
ment ; Kiitzing's Phycologia Germanica (Nordhausen,
1845, Svo). It includes the entire flora, and only appeared
a few weeks after the preceding work, Roemer's materials
having been made use of ; yet it is carried out perfectly

BOTANICAL GEOGRAPHY. 431

independently of these, and although subject to known
criticisms in a systematic point of view, is indispensable to
the understanding of the author's larger work on the Algae.
Local Floras of Germany: F. Wimmer, Flora of Silesia.
Concluding volume (Breslau, 1845, 12mo); J. C. Metsch,
Flora Hennebergica, a contribution to the Flora of the
Prussian portion of the Forest of Thuringia (Schleusing,
1845, 8vo); F. Schultz, Flora of Palatinate (Speier, 1846,
8vo) ; it appeared, however, in 1845.

In Metsch's memoir upon the plants of the mouth of
the Swine (Ratisbon Flora, 1845, pp. 705-8) is contained
a sketch of the vegetable formations on the island of
Usedom.

The sandy soil is in some places extended into plains, at others depressed,
so as to receive deposits of peat or salt-water lakes ; sometimes it forms ele-
vations, which are partly covered with the pines, or even considerable forests
of beech. The dunes along the coast are consolidated by roots of Gluma-
cese or Salix. A few of the characteristic plants only can be mentioned
here, as the author only enumerates the more rare species :

1. Formation of plants on the dunes, e. g. Ammophila arenaria and Bal-
tica, Mymus arenarim, Carex arenaria, Kochia hirsuta, Halimus portulacoides,
Petasites spurim, and Anthyllis maritima.

2. Formation of the sea plants, e. g. Aster salignus, Erythrtea linariifolia,
Zannichellia pedicellata, Juncus balticus, Scirpus Rothii, and Hierochloa
borealis.

3. Formation of the marsh plants, e. g. Thalictrum aquilegifolium, Barbarea
stricta, Helosciadium inundatum, Lysimachia thyrsiflora, Euphorbia palustris,
Salix daphnoides and rosmarinifolia, Stratiotes, Carex filiformis, and Calama-
grostis stricta.

4. Formation of the peat plants, e. g. Ledum palmtre, Betula fruticosa,
Empetrum, and Myosotis sparsiflora.

5. Formation of herbaceous plants on sunny hills, e. g. Thalictrum minus
and simplex, Silene viscosa, and Ononis hircina.

6. Formation of the forests, e. g. Arabis arenosa, Vicia villosa, Peuceda-
num Oreoselimim, Arctostaphylos officinalis, Pyrola chlorantha, media, and
umbellata, and Goody era repens.

V. Mohl has written a memoir on the Flora of Wur-
temberg (Wiirtemb. naturwissenschaftliche Jahreshefte.
Jahrg. i, pp. 69-109. Stutt., 1845, 8vo).

He commences with general remarks on the scientific importance of local

432 BOTANICAL GEOGRAPHY.

floras. He states their object to consist in the investigation of the limits
of the distribution of the species within a larger district, and for this purpose
to compare them, as e. g. the flora of Wurtemberg with that of the adjacent
countries. In this manner he shows, that when a natural subdivision of
Germany is made, the flora of "Wurtemberg belongs to those of the adjacent
districts, and does not contain distinct centres of vegetation of its own.
Mohl distinguishes four separate regions : the fluviatile system of the Neckar
and the Tauber, the Black Forest, the rugged Alp, and the tertiary plain of
Upper Swabia.

1. The Neckar region, lying between the Swabian Jura and the Black
Forest, in regard to the distribution of its plants, may be considered as a
portion of the Rhine district. The eastern limit to the occurrence of a
tolerable number of plants exists at the Jura ; but with the single exception
of Orobus albus (at Tubingen), no species is found from the Neckar to the
Tauber, which does not also exist in the valley of the Rhine. But the dis-
trict on this side is poor in comparison to the latter ; for " in correspondence
with a general phenomenon," as the bed of a river becomes narrowed, many
plants disappear which are common lower down the stream. The author
also gives a list of more than fifty species which prove this connexion, and
from which we shall select the following as characteristic forms of the Rhine
district : Helianthemum celandicum (vineale), Myagrum perfoliatum, Isatis
tinctoria, Diplotaxis tenuifolia and muralis, Altfuea hirsuta, Lathyrus hirsutus,
Rosa gallica, Helosciadium nodiflorum, (Enanthe peucedanifolia, Carum bul-
bocastanum, Crepis pulchra, Lactuca saligncf; Artemisia pontica, Centaurea
nigra, Heliotropium Europaeum, Calamintha officinalis, Mentha rotundifolia,
Parietaria diffusa, Spiranthes testivalis, and Scirpus mucronatus. Geologically
considered, the district of the Neckar and the Tauber belongs to the muschel-
kalk, lias, and the keuper formations. Of these, the muschel-kalk, as in
Thuringia, exerts the most important influence upon the distribution of the
plants, whilst the lias and keuper, being less homogeneous formations, favour
a greater chemical variation in the soil. A list of about 20 sand-plants con-
trasted with 100 plants belonging to the muschel-kalk, shows how the latter
augments the number of indigenous species to a greater extent than the other
formations.

2. The Black Forest, the soil of which is derived from bunter sandstone or
Plutonic rocks, contains in the district of Wurtemberg but few Phanero-
gamia peculiar to it, whilst the higher elevations of these mountains, which
are also poor in plants, belong to Baden. Among the plants of the Black
Forest of Wurtemberg, excepting those which also occur in other regions of
the kingdom, there is not one which is not distributed over the greater part
of the mountains of Germany, so that e. g. all those mentioned, excluding
Crocus vernus, occur also on the Hartz. If we bring into relation with these
facts Kirschleger's general remarks upon the whole of the Black Forest (see

BOTANICAL GEOGRAPHY. 433

Ann. Rep. for 1843), we cannot enumerate these mountains among the
independent centres of vegetation of the flora of Germany, because the
whole of their Phanerogamia may be regarded as having migrated from the
Alps, the Vosges, the Jura, or the Rhenish mountains.

3. The rugged Alp (Swabian Jura) possesses the characteristic vegetation
of the Jura-limestone, which is uniformly distributed from Switzerland to
Franconia. However, although the mean level of the elevated surface
amounts to more than 2000', and individual summits ascend to more than
3000', the alpine forms of plants, which are common on the higher Jura of
Switzerland, are for the most part absent here, and even the few species
(7 sp.) which belong to this category, are in most cases found at single spots
only ; whilst on the other hand, many calcareous plants from the valleys of
the Bavarian Alps are common here. About 50 species are found in "Wur-
teniberg on the rugged Alp only, 34 calcareous plants occur in common with
the region of the Neckar, 1 8 species with Upper Swabia only, 16 others
with both these districts, and 5 with the Black Forest. Perhaps the follow-
ing from the list of plants peculiar to the Swabian Jura might be mentioned
as characteristic forms, omitting those which are diffused over the calcareous
Alps : Thalictrum galioides, Thlaspi montanum, Sisymbrium austriacum, Ery-
simum crepidifolium and odorattim, Dianthus ctesius, Linum flavum (at Ulm),
Coronilla montana and vaginalis, Sorbus latifolia, Leontodon incanum, Doroni-
cum Pardalianches, Jasione perennis, Specularia hybrida, Digitalis lutea,
Nepeta nuda, Orchis pallens, Aceras anthropophora, and Iris Germanica.

4. The tertiary plain of Upper Swabia lying between the Jura and the
Alps, 1250' to 2000' above the level of the sea, is geographically a portion of
the plateau of Upper Bavaria, and also contains its vegetation, whilst the
Jura agrees with it far less than might have been expected. Even the peat-
moor formation is the same here as in the bogs of Bavaria. Upper Swabia,
although it has been least explored botanically, probably contains more plants
than any other part of Wurtemberg, on account of its fertile calcareous
Molasse-soil, the considerable variations of its elevation, the abundance of
water, and the proximity of the Alps from which, as in Bavaria, many plants
are washed down. The list of those plants of Upper Swabia which have
not hitherto been observed in other parts of Wurtemberg, includes about
90 species.

The following from among them, excluding the alpine plants, may be men-
tioned as characteristic : Ceratocephalusfalcatus, Viola lactea, Linum viscosum,
Alsine stricta, Potentilla norvegica, Saxifraga Hirculus, Helosciadium repens,
Gentiana wtricularis, Pedicularis sceptrum, Primula acaulis, Betula humilis,
Stratiotes, Iris graminea, Allium suaveolens, Juncus tennis, Carex capitata,
microglochin, chordorrhiza, cyperoides, and Heleonastes.

Mohl makes some ingenious observations on the distribution of alpine
plants towards the Bavarian plateau of Upper Swabia. He distinguishes

434 BOTANICAL GEOGRAPHY.

several kinds of distribution : 1. The seeds are constantly being carried down
anew by the waters, and the individuals which germinate are, therefore, only
accidental inhabitants of the drift on the banks, not having any fixed locality,
e. g. on the Iller. Campanula ceespitosa, Hutchinsia alpina, &c. 2. Other
alpine plants, which also grow upon the Alps themselves, on the drift of the
rivers, again meet with the conditions necessary for their existence in the
elevated plains : hence they constitute a permanent formation there, e. g.
Myricaria, Salix daphnoides, and Epilobium rosmarinifolium. 3. Other plants
of the alpine flora occur in the plain of the peat-moors, far distant from
the present alpine rivers, distributed socially, e. g. Bartsia alpina, Primula
auricula, Gentiana acaulis, in large masses on the bogs of Upper Bavaria,
and Veratrum album also in Upper Swabia. On the Alps, part of these plants
grow in totally different localities; yet, according to the opinion of the
author, there is no doubt that, like those above mentioned, they emanated
from the Alps, although the conditions under which these depositions occurred
cannot now be ascertained. In this respect, he declares that Zuccarini's
view is a very hazardous hypothesis, who supposes that the first seeds were
carried down in remote ages by the same rivers which filled up the whole of
the tertiary plain with alpine Molasse, and gave rise to the continent. This
view is inadmissible, because the phenomenon of the occurrence of the alpine
plants in the peat-moors is evidently the same as that which is now going on
in the north of Germany, where e. g. Primula farinosa, Swertia perennis, and
Salix daphnoides are met with under the same conditions. The humous
pasture-soil of the Alps is not so entirely different from peat, nor the climate
of Upper Bavaria so very different from that of Mecklenberg, as regards
many plants, as to render inadmissible every explanation of this simultaneous
growth of individual species in remote plains and on the mountains, by means
of the soil and climate : we need not then hypothetically devise any geolo-
gical causes. Are not aerial currents sufficient to convey the minute seeds
of the Gentianese, or the cotton of a willow, to all those parts of Germany,
nay, even of Europe, where the climate and soil permit their germination
and growth ? What their original locality was, whether a plain or a moun-
tain, appears to me an idle question, because it is incapable of scientific
solution. 4. The same applies to all those alpine plants which have attained
any extent of diffusion in the south-east corner of Upper Swabia, e. g. Rho-
dodendron ferrugineum, Campanula barbata, Streptopus amplexi/olius, &c.
Mohl considers that these are the original plants of Upper Swabia, and that
their origin is not to be looked for in the Alps ; this view will appear totally
untenable to every one who is acquainted with the botanical relations of the
Alps, from personal observation, although we are not in a condition to ascer-
tain how they arrived at their present locality. The latter point appears
simple to me, when we recollect that the greater number of these plants
thrive also on the Sudeten and other remote mountains, and thus probably

BOTANICAL GEOGRAPHY. 435

possess a wide climatic sphere, and also means of more easy diffusion by the
air ; but how the first question can be decided by personal observation, I do
not understand, inasmuch as a plant may be diffused equally as luxuriantly
and generally on a secondary as on a primary locality, as e. g. the thistle of
the Pampas of Buenos Ayres teaches us, which in the Old World, where it is
indigenous, is found at individual spots only, whilst in the former place it
covers the plains in the most intimate community. The conclusion of this
important work is formed by a catalogue of the names of all the Phaneroga-
mia hitherto found in Wurtemberg, without the localities ; it contains only
1287 species, i. e. more than 100 species less than are known (according to
my manuscript) in the kingdom of Hanover : hence Mohl appears to be jus-
tified in the opinion, that many remain to be discovered in Wurtemberg.

The Topography of the Upper Pinzgau (see Ann. Rep.
for 1840), contains a work by A. Sauter, with which
I am only at present acquainted from Beilschmied's
Abstract (Ratisbon Flora, 1845, pp. 501-7), upon the
botanico-geographical relations of this district, which in-
cludes the longitudinal valley of the upper Salzach, between
the Tauern and the clay-slate alps of Kitzbiihl.

It contains, in addition to lists of the more rare species, a sketch of the
botanical regions, but the source of the altitudes given is not stated. 1. Region
of the cultivated country. 2400' 4000' on the south side, 3000' on the
north side of the mountain. Pasture land alternates there with forests ;
meadows and fields are more rare. Most of the deciduous trees, and Alnus
imana is very common, do not ascend higher. 2. Forest region. On the
average 3500' 5500'. Finns abies, however, which constitutes the main
part, appears only to thrive as high as 5000', P. Picea at 4000', whilst
P. Cembra here and there covers the upper slopes, and in the Tauern chain
ascends as high as 6000', the same height as P. Larix. 3. Alpine Region.
On the average, 5503' 8000'. It also contains but few meadow surfaces,
mostly naked rocks and detritus. The sub-alpine forests do not form a
dense zone there; Rhododendron ferrugineum occurs in groups as far as
6000'; dwarf willows, Empetrum, Arctostaphylos, and Azalea procumbens, as
far as 7000'.

Perini read a paper on the Botanical Regions of Trient,
in the south of Tyrol, before the Association of Italian
Naturalists (Atti di VI riunione, p. 460).

L. v. Heufler has described a botanical excursion in
the north of Istria (Die Golazberge in der Tschitscherei.
Triest, 1845, 4to).

436 BOTANICAL GEOGRAPHY.

On the 16th of June the author collected 300 species of plants on a
mountain-ridge of the Karst, only 3410' in height, to the south of the Fium-
mean road ; they are enumerated in the order of their occurrence in this
luxuriously-printed work. In the map which is appended, the following
regions of the coast-country of Illyria, in the direction from the Adriatic Sea
to the coast of Terglou, are distinguished, but how the altitudes were deter-
mined is not stated : 1, 0' 500' ; region of Olives. 2, 2000' ; oak-region
(with regard to which it is incorrect to state that the region of the species
of the north of Europe is not different from that of the Mediterranean species).
3, 2000' 4800'; beech region. 4, 6500' ; pine region. 5, 8500'; region
of the alpine plants. 6, 9036' ; region of snow. The vegetation of the
Golaz mountains resolves itself into the oak forests (1500' 2000' : Quercus
Robur, pedunculate, Cerris, ^.n^pubescens}, beech forests (2000' 3410'), alpine
pastures, and the rocky formation. In addition to this main subdivision,
separate arrangements in groups are also mentioned, e. g. shrubs of Ornus
in the lower, and of Corylus Avettana in the upper part of the oak-region,
herbaceous meadows of Cyiisus and Genista, &c,

Wierzbicki has published a Catalogue of the Plants
found in the Banat (Ratisbon Flora, 1845, pp. 321-25),
subsequent to the appearance of the most recent work on
the flora of this province (Rochel's Travels in the Banat,
1838); also, Prof. Fuss, of Hermannstadt, a list of 319
plants of Siebenburg, with their localities (Archiv des
Vereins fur Siebenburg. Landeskunde, (Bd. 2, Hft. 3).

O. Heer's memoir upon the Upper Limits of Animal
and Vegetable Life in the Swiss Alps (Zurich, 1845, 4to),
is more important in a geological than a botanical point of
view, on account of the descriptions and illustrations of
new insects belonging to the snow-region contained in it ;
however, it also contains some valuable observations on
the forms of plants, which, under certain conditions,
vegetate above the snow limit (8500').

Lichens extend far above the Phanerogamia and Mosses ; they exist even
on the summit of Mont Blanc. The highest of the Phanerogamia was
Androsace glacialis (pennina Gaud.), and occurred at 10,700' on Pic Linard ;
and from this altitude down to 10,000', the following were found in succes-
sion on different glaciers, i. e. on account of the position or inclination of
the surfaces free from snow on the Rhetian Alps : Oentiana bavarica,
var. imbricata, Silene acaulis, Chrysanthemum alpinum, Ranunculus glacialis,
Cerastium latifolium, var. glaciale, Saxifraga oppositifolia and bryoides, Cher-

BOTANICAL GEOGRAPHY. 437

leria and Poa laxa. Associated with these between 10,000' and 9000' we find
50 more, and as far as 8500', i. e. at the snow-line, 46 others; so that the
entire flora of the Rhetian Alps consists of 106 Phanerogamia, belonging to
23 families. All these plants are perennials, most of them casspitose,
hence they are propagated without the seed arriving at maturity ; all of them
are depressed and small, and are thus less influenced by the heat of the air
than of the soil : in fact, the only two woody plants are dwarf willows, their
stems being almost entirely inserted into the earth. Yet the temperature
of the soil at these altitudes is probably for a short time only above the
freezing point. The author very correctly explains their being enabled to
grow, notwithstanding this circumstance, by the short period of their vege-
tation, as when transplanted into the low. country, they become, without
exception, spring plants, which, in a few weeks after budding, ripen their
fruit, their winter sleep being proportionately prolonged. Moreover, in the
low country they all exhibit but little susceptibility to cold, so that even at
the period of their flowering, although exposed to frost, they are not at all
injured. Even if, in their elevated position, the spring season should not occur,
they would endure a state of hybernation for several years, without being
destroyed. The conditions of vegetation being so different from those of the
level country, explains the fact that the Phanerogamia of the snow-region
never become spontaneously distributed in the valleys. The case is different
with the Cryptogamia ; for the lower the degree of organization, says Heer,
the less does the form require to be modified, to adapt it to a different
climate.

Mougeot and Nestler have published, in connexion with
W. P. Schimper, the twelfth century of their well-known
collection of dried Cryptogamia from the Vosges (Stirpes
Cryptogamae Vogeso-Rhenanae, fasc. 12. Bruyere, 1844,
4 to).

French local floras and systematic contributions upon
French plants : Observations sur quelques Plantes Lor-
raines par Godron (Nancy, 1835, 8vo, pp. 31), con-
taining a supplement to his Flora of Lorraine; Choulette,
Synopsis de la Flore de Lorraine et d'Alsace, Partie i,
Tableau Analytiques (Strasb., 1845, 16mo); Cosson et
Germain, Flore descriptive et analytique des Environs de
Paris (Paris,1845, 8vo, 2 vols.), remarkable for its accuracy,
and the systematic investigations contained in it ; it gives
satisfactory elucidations, e. g. of Astrocarpm Clusii, Tri-
folium Parisicme, Euphrasia Jaubertiana, Potamogeton
tuber culatus, tmdCarevMairii; Puol, Catalogue des Plantes

438 BOTANICAL GEOGRAPHY.

qui croissent dans le Departement de Lot (in the Annuaire
du Departement, pp. 1845, 1846), extending to Hexan-
dria ; F. Schultz, Continuation of the communications
on the Orobancheae of France (Ratisbon Flora, 1845,
p. 738) ; Desmazieres, Eleventh Contribution to the
knowledge of the Cryptogamia of France, containing
Fungi (Ann. Sc. nat., 1845, 3, pp. 357-70).

The excellent observations of Ch. Martins on the cli-
mate of France, which were mentioned in the preceding
Annual Report, are now. published in greater detail, and
have been augmented by a description of the botanico-
geographical relations (Fssai sur la Meteorologie et la
Geographic botanique de la France ; separate division of
the encyclopaedic work, Patria; La France ancienne et
moderne. Paris, 8vo).

The French botanical geography, however, is merely based upon the facts
given in Duby's ' Botanicon Gallicum.' The distribution of about 3700
Phanerogamous plants through France is shown by a series of lists. 1.
1250 species are diffused over the entire country; i. e. they occur in the
local floras of Boreau, Godron, Cosson and Germain, and Dumortier, and in
Bentham's ' Catalogue of the Mora of the Pyrenees.' 2. About 30 species,
most of which are distributed over Central Europe, correspond, in France, to
the climate of the Vosges and Seine. (See the preceding Ann. Rep.) 3.
About 30 species belonging to the valley of the Rhine, are confined to the
Vosges climate : the mountain plants of the Vosges, which, however, appear
also to occur on other mountains of France, are separated from these, as
also the southern forms of the valley of the Rhine (10 species), but which
according to their distribution ought rather to be arranged in the third list.
4. About 30 species of plants belonging to the north-west, corresponding to
the climate of the Seine. 5. The centre of France forms a transition-
district from the north to the south, and has only 3 species peculiar to it.

6. 750 species belonging to the south of France, correspond to the climate
of the Garonne and Rhone, but they also occur in the Mediterranean district.

7. 800 species are confined to the Mediterranean climate. 8. 500 species
belong to the subalpine region of the mountains of France, the level of which,
between the 46 and 49 of north latitude, Martins estimates at from 600 m.
to 1600m.; south of 45, from 1000m. to 1800m. 9. 300 species grow
beyond these limits in the alpine region. The conclusion consists of
plants arranged according to the localities.

In the same paper Martins also publishes the following measurements of
the limits of vegetation in Dauphiny :

BOTANICAL GEOGRAPHY. 439

Cultivation of barley. Upper limit.

Col de la Vachere. North side, 1745 m. South side, 2110 m.
Fagus sylvatica. Upper limit.

Grande Chartreuse, 1465 m. Col des Sept Lacs, 1 475 m.
Pinus abies. Upper limit.

Grande Chartreuse, 1631 m. In a shrubby form, 1900 m.
P. picea. Upper limit.

Col des Sept Lacs. North side, 1770 m. South side, 2045 m.
Alnus viridis. Upper limit.

Col des Sept Lacs. North side, 1910 m.
Sorbus aucuparia. Upper limit.

Col de la Vachere. North side, 2000 m.
Rhododendron. Lower limit.

Col des Sept Lacs. North side, 11 60m. Grande Chartreuse, 1660m.
Col de la Vachere. North side, 2125 m.
Upper limit.

Col de la Vachere, 2410m.
Pinus Cembra. Lower limit.

Coldela Vachere, 1740m.
Upper limit.

Col Longet, 2515 m.
P. larix. Upper limit.

Col Longet, 2515 m.

Daum has described two barren, almost desert regions on
the south coast of France (Bemerkungen liber die Land-
wirthschaft in Siidfrankreich. Charlottenb. 1844, 8vo).

The plain of Crau, which lies to the south of Aries, covers nearly seventy-
five square miles ; it consists of a gravelly surface, containing scattered,
but nutritive herbs and grasses, upon which no kind of agriculture can be
carried on, but 30,000 fine sheep find pasture from late in the autumn until
the spring, when they are driven upon the pastures of the Maritime Alps ;
and which it is now being commenced to convert into meadow-land by arti-
ficial irrigation ; next the plain of Camargue, in the delta of the Rhone, a
boggy salt-marsh, nearly half of which consists of flooded land and bog, the
remainder of pasture-land and a few fields, and which, by means of a large
capital, it is also being attempted to turn to advantage by canal drainage.
As regards the agriculture of the province, the traveller remarks that, on
account of the grape-vine requiring a large quantity of manure, without
affording nutriment to cattle, the principal object is the cultivation of fodder,
for as there are no meadows, this is necessarily obtained from lucerne.
These facts show the natural character of the country.

Cuynat's Topography of Catalonia (Memoires de

440 BOTANICAL GEOGRAPHY.

F Academic de Dijon, 1845), contains a Catalogue of the
Plants found in this Spanish province (2, pp. .91-100).

Six hundred species are enumerated, but most of them are so much more
extensively distributed on the Mediterranean coasts, as to prevent our ob-
taining an accurate idea of the peculiar nature of the vegetation of Catalonia,
which has not yet been described.

Willkomm's sketch of Monserrat, with which he con-
cludes his Botanical Reports upon Spain, may be men-
tioned as a contribution towards filling up this deficiency
(Bot. Zeit., 1846).

This isolated conglomerate mountain, which the traveller visited in April,
is scarcely more than 3000' in height ; but the summit is only accessible by
a deep rocky valley, which runs in a north-westerly direction, whilst the
outer sides rise so steeply that they cannot be ascended. In Catalonia,
the " warm region," which undoubtedly corresponds to the region of Cha-
mcerops (see below), extends scarcely more than 1000' ; hence the greater
part of the Monserrat belongs to " the mountainous region" (the region of
evergreen oaks), and the summit reaches the subalpine region (region of the
pines). The Mediterranean, as also the Central European plants, are mixed
on this mountain with a number of Pyrenean plants. In the lower region,
the heights at Bruch are covered with forests of Pinm halepensis and pinea ;
the other parts are covered with freely-vegetating " montebaxo," consisting
of evergreen oaks, Pis facia lentiscus, Erica arbor ea, and other shrubs.
Characteristic plants : Genista hispanica, E^tphorbia oleifolia G., Globularia
Alypum, Cor is monspeliensis, and Passerina tinctoria Pourr. At the central
elevation : Poly gala saxatilis Lag. ; Erodium supracanum, Sarcocapnos ennea-
phylla, Carduus tenuiftorus Salzm. ; Ramondia pyrenaica, and Convolvulus
saxatilis. The upper region was not developed at that season ; however,
Arctostaphylos uva ursi, Globularia nana Lam., and Narcissus JonquUla, were
in flower.

The families containing most species in the flora of Castile form the
following series, according to Renter's collection, which contains 1232 sp.
(Boissier, Voy. en Espagne, i, p. 207) : Graminacese (161 sp.), Leguminosse
(130), Synantheracese (125), Cruciferse (74), Caryophyllaceae (52), Rosacese
(38), Ranunculacese (33), Boraginese (31), and Chenopodeacese (26).

According to Willkomm (loc. cit.), the Sierra Morena contains an uncom-
monly uniform vegetation, notwithstanding its great length and breadth.
With an average breadth of 8 geogr. miles, it extends from Murcia to Algarve,
only forming, however, an intermediate mountain chain, the crest of which,
for the most part, only ascends to 2 3000', and the greatest elevations of
which are hardly 5000'. By the density of its forests or tall shadowing

BOTANICAL GEOGRAPHY. 441

shrubs, and by this connected green and fresh vegetation, the Sierra Morena
differs from all other mountains of Andalusia, which only contain isolated
spots of wood, and a low, barren " montebaxo." Geologically considered,
the principal rocks of the Sierra Morena consist of sandstone, which, in the
form of grauwacke, forms the greater part of the mountain-chain, occurring
from 4 to 6 geog. miles broad, in gently-rounded mountains and undulating
summits, alternating at Almaden with clay slate, in the province of Huelva
with gneiss, and southwards, near the low plain of the Guadalquiver, inclosed
by other sandstone formations. The central portion of the mountains is
interrupted by the immense granitic formation of Cordova, which, in the
form of an elevated plain, inclines from Hinojosa towards the north, and
becomes connected there with white quartzose rocks, which, between Almaden
and Fuencaliente, appear to form the highest chain of the whole Sierra.
In the opinion of the traveller, the character of the vegetation is associated
with these geological variations, which are lithologically unimportant. The
predominating shrub on the grauwacke, as far as Portugal, is Cistus ladini-
ferm, which extends over the Sierra Morena for a length of more than 50 geog.
miles, and " frequently covers whole square leagues exclusively." Next to
this, the most common are Phillyrea angustifolia, Rosmarinus, and a Heli-
antliemum. The forests on the grauwacke consist of evergreen oaks, of
Quercus Ilex, Ballota, and Suber, but the first is mostly only a shrub ; the
dry, arid, but densely-populated granite plateau is very sterile, but still pos-
sesses extensive woods of Quercus Ilex and Ballota, only a dwarf growth,
however, of Quercus Ilex, mixed with Cistus ladaniferus, PMllyrea angusti-
folia, and Arbutus unedo. The southern sandstone chains are furnished with
an extremely luxuriant and varied " montebaxo," which, near the city of
Cordova, alternates with woods of pines and cork-oaks. The quartzose rocks
of Mancha are also covered by a "montebaxo" abounding in forms, among
the shrubs of which Cistus populifolim is distinguished. Finally, in Huelva,
Portuguese forms are associated with the other shrubs, as Genista tridentata,
Ulex genistoides, and with these Erica umbellata, Teucrium fruticans, and
Helianthemum halimifolium. Unfortunately, Willkomm has not made us ac-
acquainted with the vernal vegetation of the Sierra Morena, which would be
the most interesting. But the dryness of the summer commences here in
July, and from that time until the autumn no more flowering plants are met
with. Different kinds of bulbous plants appear very uniformly distributed
in the autumn, as Scilla maritima, Scilla autumnalis, Leucojum autumnale,
Merendera Bulbocodium, &c.

Boissier's large work upon the southern border of
Granada and the Sierra Nevada forms the most valuable
contribution to botanical geography which has been made
during the past year (Voyage botanique dans le Midi de
de 1'Espagne, t. i. Narration and Geographic botanique,

442 BOTANICAL GEOGRAPHY.

pp. 241. Paris, 1845, 4 to) ; as regards the earlier sys-
tematic parts of this illustrated work (t. ii), see the Ann.
Rep. for 1840-1.

JBoissier's excellent account includes the coast-terraces between Gibraltar
and Almeria, towards the centre of the country as far as the elevated sur-
faces of Andalusia, and thus entirely includes the highest mountain-chains
of the south of Spain. Along the entire coast-line a series of isolated moun-
tain-chains, consisting of marmoraceous limestone, arise directly, in almost
every case, without any intervening land, the western extremity always
ascending highest, whilst towards the east the ridge gradually falls : to this
system belong the Serrania de Honda (6000') 3 Sierra Tejeda (6600'), and
Sierra Gador (7000'). These chains, which run parallel with the coast, may
be considered as forming the southern mountain-border of the Spanish pla-
teau ; for its northern foot, at an elevation of from 2000' to 2500', passes
directly into the elevated plain of Honda, the Vega in Granada, or the great
plains of Guadix and Baza. In a line from Ducal to Almeria, not far from
Granada, the chain of the Sierra Nevada, twenty-two leagues only in length,
is inserted between the boundary chain and the elevated plain ; it is nearly
twice their altitude, but narrow, its highest summits ascending to an elevation
of 11,000'. In fact, the passes in the western portion are not situated below
9500', whilst toward the east the mean height of the crest appears to dimi-
nish to 6000'. The Sierra Nevada is mainly composed of mica-slate, but on
its flanks secondary and tertiary formations have been carried up with it as
far as an altitude of 6000'. The district of Alpujarra forms an important
constituent of these mountains ; it includes the longitudinal valley running
between the coast-chain S. Contraviesa and the Sierra Nevada, together with
the southern, intersecting valleys of the latter. The following are some
of the heights which were measured by Boissier with the barometer : The
city of Honda, on the plateau, 2300'; Granada, 2200'; Sierra Nevada, the
farm of San Geronimo, 5064'; Col de Vacares, 9472'; Picacho de Veleta,
10,728'; Mulahacen, 10,980'.

Four botanical districts, which Boissier distinguished in South Granada,
yielded him vascular 1900 plants, which he is inclined to regard as forming
three fourths of all the indigenous plants of this district. The author con-
siders the following as among the general characters of the flora viz. that
many forms cover the soil in gregarious condition, and that the south of
Spain contains more thorny plants than any other country in Europe, and
hence resembles the steppes of Western Asia, although the families which de-
velope the thorns are not the same. The hot region (region chaude) comprises
the coast declivity up to a level of 2000'. Intense atmospheric precipitations
fall during October and November ; the vernal rainy season, which is less
regular, lasts from February to March, sometimes until April ; uninterrupted
drought prevails from April to the end of September. Thus the dry season

BOTANICAL GEOGRAPHY. 443

there, is probably of longer duration than in any other point of the Mediter-
ranean flora. Observations are given, made at Malaga, during nearly three
years (1836 1839), by Haenseler, on the distribution of heat; the extreme
temperatures of which, as also the monthly average, calculated from the
corresponding months of the year in which the observations were made,*
yields the following values :

Med. Max. Min.

January . . 12'25 17'22 6'2

February . . 14 "3 18 "25 6 '1

March ... 15 -8 21 '62 10 -0

April ... 17 '8 25 '0 11 '25

May ... 21 '2 24 -5 15 '72

June ... 23 -4 26 -87 20 "12

July ... 26 -2 31 '87 23 '5

August ... 26 -8 30 '6 23 "75

September . . 24 '4 29 '87 19 -37

October . . 22 "25 25 '5 19 '25

November . . 18 "15 22 "75 11 '2

December . . 15 '75 21 '0 8 '5

Mean annual temperature, 17'3

The vegetation passes through phases corresponding to this climate.
After the dry season, Liliacese are developed during the first rain of October
or November ; these are succeeded by the annuals, which flower throughout
the entire winter. The flowering season of most of the plants is in April
and May; in June and July, when all the annual plants have withered, her-
baceous plants belonging to the families of the Synautheracae, Umbelliferae,
and Labiatse flower ; lastly, from August to September, the most profound
repose of vegetable life prevails, so that two or three Liliacese, Mandragora,
and Atractylis gummifera, are all that remain. The hot region is principally
characterised botanically by Chamaerops, which covers large tracts and pre-
vents cultivation ; as in Valencia, it only ascends to 2000'. Among the
cultivated plants, the orange also corresponds accurately to the extent of this
region. The soil of other parts is principally devoted to the cultivation
of the grape-vine, the fruit of which ripens at the end of August. The
Cerealia require artificial irrigation : on those parts which are reached by the
water from the mountains, either in its natural descent or by aqueducts, we
sometimes find most luxuriant fields of maize and wheat, shaded by orange
and mulberry trees. But such oases are rare on these bare and arid slopes,
on which wheat is reaped as early as the latter half of June, and barley in
May. However, on a narrow coast-district which surrounds the coast-chains,

* The average which I have calculated refers, for June, July, and August,
to two, and for the other months to three years.

444 BOTANICAL GEOGRAPHY.

sometimes in the form of a surface containing salt-water lagunes, at others
as a line of hills, and which at Malaga alone is covered with a more exten-
sive alluvial plain, the cultivated plants are confined to the hot zone (0' 600'),
as the sugar-cane, the cotton-plant, and sweet potato, the Date-palm and
Ceratonia, as also the migrated Agaves and Opuntias, with several indigenous
plants, as Aloe perfoliata and Withania ; excepting the white poplar,
indigenous trees are absent from this littoral district. Boissier enumerates
altogether 19 trees as belonging to the hot region, part of which, however, like
the Agrumae, are of foreign origin. The following only can be regarded as in-
digenous: Ceratonia, which ascends 2000', Zizyphus, Punica, Celtis australis,
and Populus alba, with those which extend into the following region, where
they become more common viz. Ficus carica (0' 3000', on the southern
slope 4000'), Olea europtea (vid. supra), Quercus Ballota and lusitanica
(3000'), Q. Suber (4000'), Q. Ilex (4500'), and Pirns Pinaster (vid. supra),
The following are the most important formations of the hot regions : a.
Maquis (Montebaxo). Shrubs from 3' 6' in height cover the greater
part of the sloping soil, consisting of Chamtfrops, several Cisti, viz. C.
ladaniferus, albidus, and Clusii ; Pistacia lentiscus, Rhamnus lycioides, and
Phillyrea, numerous Genista, most commonly Genista umbellata and Retama
spheerocarpa, and some oaks, beneath which numerous annuals and Grasses
flower in the whiter and spring, and, more rarely, herbs which are developed
at a later period. Shrubs of Nerium denote the humid soil of the banks of
rivers, b. Campi. On bare wastes are found predominating Thymbra capi-
tata, Lavandula multifida, Teucrinm Polium, and numerous herbs, among
which Centrophyllum arborescens is pre-eminent. In other places, these are
replaced by the social Macrochloa tenacissima. In addition to these two
principal formations, there are the Halophytes of the littoral district, those
plants which are indigenous to the marshes of Malaga, and lastly, the plants
of the cultivated part of the country, with its hedges of Agaves and Opun-
tias. The following plants belong to the endemic forms of the hot region
of Granada : Caltha europcea (Celastrus Voy^], Genista umbellata and
gibraltarica, Sarothamnus beeticus and malacitanus, Ulex Ixsticus, Leobordea
lupinifolia, Ononis Gibraltarica and filicaulis, Maeoselinum, Lagascce and
fcetidum, Lonicera canescens, Withania frutescens, Triguera ambrosiaca, Lycium
intricatum, Lafuentea rotundifolia (according to Willkomm, but it is absent
according to Boissier), Digitalis laciniata, Sideritis lasiantha and arborescens,
Salsola Webbii, Passerina canescens and villosa, Osiris quadrifida, Euphorbia
and trinervia, Quercus Mesto, Salix pedicellata, and Ephedra

The second region (region montagneuse), or the region of the Spanish
plateau, is peculiar to Spain, and cannot be compared with the mountain-
vegetation of other European countries. By way of introduction to Boissier's
description, I shall premise here a remark upon the climatal cause of this

BOTANICAL GEOGRAPHY. 445

peculiar condition. In Italy, Dalmatia, and in Turkey, we find immediately
above the evergreen region, slopes abounding in forests of Central European
forms of trees, and other plants indigenous to this side of the Alps : angiosper-
mous trees, which lose their leaves in winter, even at an altitude of 1200' to
1500', frequently begin to denote this second Central European region. In
Spain, Boissier, with other authors, distinguishes two evergreen regions : a lower
one, which in the character of its vegetation appears to agree with the Italian
or Dalmatian, and reaches to 1500' in Catalonia, and to 2000' in Granada ; and
an upper one, which, extending from 2000' to above 5000', includes the greater
part of Spain, and has no analogy with any other throughout the entire south
of Europe. It has been shown by Schouw's investigations, that the climatai
cause of the evergreen vegetation of the Mediterranean lies in the aridity of
the summer, to which the plants of the north of Europe are not subjected.
Out of Spain, the latter again meet with conditions necessary for their
existence on the mountain-chains of the south of Europe, in the vicinity of
the region of clouds, where, even in the summer, the air contains mists formed
from its watery vapour, and where the low scale of temperature is the same
as in the climate of the north. The elevated plains of Spain are, however,
in summer even more arid than the coast-regions ; the humid and mild spring,
which stimulates all the plants to flower, is there succeeded by a hot and dry
summer and a cold winter ; the three seasons of the steppes of Russia are dis-
tinguished. Although this explains the fact that some plants of the plateau
of Spain recur in the Crimea, or on the elevated surfaces of Asia Minor, yet
their number is small ; for the contrast of the insular and the extreme climate
of the interior of the continent exerts such influence here, that the greater
part of the plants of Spain are not exposed to the great winter-cold of the
eastern elevated surfaces and steppes. Hence the greater part of the flora
of the plateau of Spain must consist of endemic plants, because these climatai
conditions do not exist elsewhere in Europe. This is still more strikingly
perceptible in central Spain (see the Ann. Rep. for 1843) than in Granada,
where the plateau-character is less developed on the mountain-slopes, and
the vegetation contains fewer forms. But it is clear that more plants of the
evergreen coast-region may occur in a climate of this kind than in that of
Northern and Central Europe. Returning to the observations of Boissier, he
estimates the region which corresponds in Granada to the plateau of Spain as
extending from 2000' to 4500' on the northern, and to 5000' on the southern
slopes. Within this region, but not far from its lower boundary, the cities
of Granada and Ronda are situated, where, in the winter, the thermometer
regularly falls 6 8 below 32 F. for a few days. At its upper limit, e. g. in
the village of Trevelez, in the Alpuj arras, the snow lies on the ground for four
months, from December to April. The summer-heat is frequently greater at
Granada than on the coast, but the nocturnal cooling is very considerable.
The distribution of the atmospheric precipitations is the same as in the lower

446 BOTANICAL GEOGRAPHY.

region, except that in the summer thunderstorms are common on the Sierra
Nevada, and hence the soil rarely dries up so completely as lower down.
The agriculture consists principally in the cultivation of wheat and maize,
the upper limit to which coincides with the boundary of the region. The
wheat is reaped in July, or, in more elevated localities, at the commencement
of August. The cultivation of fruit trees extends to the same level as that
of wheat ; the chesnut, mulberry, and walnut to 5000' ; pears and cherries
somewhat higher (the latter, in parts, to 6500'). But the most remarkable
phenomenon is, that here, quite independently of their horizontal area, the
olive and grape-vine extend to nearly the same level (Oka on the northern
slope to 3000', on the southern slope to 4200' ; Vitis to 3500' and 4200').
The formations of the second region are nearly the same as in Castile : a.
" Maquis" of the same aspect as in the lower region, but composed of mostly
different species. Genistese and Cistese are more common here ; those most
so are Cistus populifolius, Genista hirsuta, with Sarothamnus arboreus, JJlex
promncialis, Daphne Gnidium, Rosmarinus, &c. b. Thin forests of Pinus
Pinaster (1200' to 4000') and P. halepensis (2000' to 3000'), or of evergreen
oaks, as Quercus Ilex, Ballota and Suber (vid. sup.) The underwood here
also consists of shrubs of Cistus, the density in the growth of which increases
in proportion as the intervals between the trees augment. Characteristic
forms of the forest-vegetation : Cistus laurifolius, populifolius, and salvifolius,
Lithospermum prostratum, Herniaria incana, Scabiosa tomentosa, &c. In the
Serrania de Honda, this thicket is replaced by a mixed kind of forest, con-
sisting of Abies Pinsapo B. (3500' to 6000'), and Quercus alpestris B. (3000 to
6000'). In addition to those above mentioned, the following are the only
other trees which occur in this region : Fraxinus excelsior (3000' to 5000'),
Ulmus campestris (2000' to 4000'), Populus nigra (2000' to 5000'), and Pinus
pinea (3000'). c. " Tomillares." Low shrubs and herbaceous plants belong-
ing to the families of the Labiatse, Synantheracea?, and Cistinese form a
dense expanse of vegetation, among which stellate patches of high turf, con-
sisting QiStipa, are distinguished. Characteristic forms : Thymus Mastichina,
zygis, and hirtus, Salvia Hispanorum, Teucrium capitatum, Sideritis hirsuta,
Helianthemum hirtum, Stipa Lagasca, Linum suffmticosum, Artemisia Bar-
retieri and campestris, Lavandula Spica and Stcechas, Helichrysum serotinum,
Santolina rosmarinifolia. d. Meadows of rigid, tall grasses, which are but
little touched by cattle, and consist of Avena filifolia and bromoides, Festuca
granatemis and Macrochloa tenacissima, cover particular slopes, e. Vegeta-
tion, consisting of Cynaracese, on the untilled fields, on the clay. f. Gypsum-
vegetation with Halophytes (See Renter's description of Castile, in the
Ann. Rep. for 1843), principally distributed over the elevated surfaces of
Guadix and Baza. Characteristic plants, mostly glaucous, and part furnished
with fleshy leaves : Peganum, Frankenia thymifolia and corymbosa, Lepidium
subulatum, Ononis crassifolia, Helianthemum squamatum, Statice, Abriplex,

BOTANICAL GEOGRAPHY. 447

Salsola, and Juncus acutus. The following are among the endemic forms of
the second region of Granada, in addition to those already mentioned, e. g.
Aplectrocapnos bcetica, Crambe filiformis, Hypericum bteticum and caprifolmm t
Rhammts velntinus, Ulex baeticus, Genista biflora and Haenseleri, Sarothamnm
affinis, Ononis speciosa, Anthyllis tejedensis, Saxifraga gemmulosa, Mtsoselinum
millefolium, Lonicera splendida, Santolina canescens, viscosa, and pectinata,
Centaurea acaulis, Clementei, prolongi, and granatensis, Cynara alba, Cha-
mapeuce hispanica, Digitalis laciniata, Salvia candelabrum, Thymus longi-
florus, Teucrium fragile and Eaenseleri, Salsola Webbii and genistoides,
Euphorbia Clementei and leucotricha, and Oligomeris glaucescens.

Third region (Bossier's alpine region), from 4500' (5000 X ) to 8000'. The
name applied to this region is not fortunately selected, because at the most
it could only be compared with that of the subalpine vegetation of the
Alps. But considering that, in addition to many endemic plants, at least a
large proportion (two fifths) consist of the plants of Central Europe, it would
have been more appropriate to have named it after them. But the question
of whether the region possesses natural boundaries is of more importance
than that regarding the name. On this point it is at once evident, that the
tree-limit, which in most mountains so sharply defines the Central European
from the alpine region, coincides in the present instance with the latter
(6000' to 7000') according to Boissier's estimation. Among the trees which
vegetate here we have Pinus sylvestris, Taxus, Saliao caprea, and Sorbus
Aria ; hence, in fact, forms belonging to the forests of Central Europe.
Now if, as in the Sierra Nevada, the forests had diminished to such an extent,
or in the course of time had disappeared, so as at the present day not to have
any considerable influence upon the natural character of the mountains, still,
to allow of comparison with other mountains, it is requisite to determine the
regions according to the sphere of distribution of those species which inhabit
a large portion of Europe, and thus afford the most certain standard for the
climate of any particular region. In the distribution of a mountain into
regions, the height at which the vegetation undergoes a decided change is
not the only point to be taken into consideration, but also where the climates
corresponding to those of other latitudes approximatively occur ; a determi-
nation which can only be made by the comparison of the vertical distribution
of the same plants. There is another decided reason, which renders it
essential to form the regions of the Sierra Nevada beyond the tree-limit (part
of Boissier's alpine region and his snow-region) into one. Boissier does not
give any other decided difference between the two, than that in the snow-
region flakes of snow remain during the summer, and that the taller shrubs
are absent. That as the altitude increases, the alpine plants themselves
alter considerably, is always found to be the case in the upper mountainous
regions ; hence the latter might be subdivided, without the accuracy and
distinctness of the representation being increased. But the region of

448 BOTANICAL GEOGRAPHY.

Genista aspalafhoides evidently corresponds to the Rhododendra of the Alps,
the dwarf birch and the shrubby willows of the north ; formations which are
always considered as belonging to the alpiue region, or have been separated
from it as subalpine. Hence I propose the following regions for Granada,
which are comparable with those of other mountainous countries in the south
of Europe, and whereby the most remarkable circumstance, that the alpine
region has a very wide, and the central European a very narrow altitudinal
extent, vanishes when brought into relation with the general fact which I
have elsewhere determined, that in Europe the tree-limit does not ascend
southward of the Alps.

A. Evergreen region. 0' 5000' (4500').

a. Region of Chamarops. 0' 2000'. (Boissier's hot region.)

b. Region of the Cisti. 2' 5000' (4500'). (Boissier's mountainous
region.)

B. Central European region, or pine-region. 5000' (4500') to 6500'. (Part
of Boissier's alpine region.)

c. Alpine region. 6500' 11,000'.

a. Region of alpine shrubs. 6500' 8000'. (Part of Alpine region.)

b. Region of alpine shrubs and grasses. 8000' ll'OOO. (Boissier's
snow-region).

But we must now follow Boissier's further description, and hence take in
the two regions which are contained in his alpine region. In the upper
part of it the snow settles even at the end of September, and the last masses
of snow do not melt until the commencement of June (hence corresponding
with the alpine region of the Alps), whilst in the lower part, the soil is only
covered with snow for four months (hence the climate of the Coniferous
region). The distribution of heat corresponds generally to the position of
the coasts ; the winter is not cold, and the temperature of the summer never
exceeds 77 E. The atmospheric precipitations are distributed over the
whole year ; in spring, and even throughout the summer, mists and thunder-
storms keep the soil moist, and this to a greater extent on the northern than
on the southern slope, which explains the greater abundance of plants on
the north side of the mountains ; consequently all the vegetative conditions of
that portion of Europe on this side of the Alps are present. Agriculture is
carried on there only as in gardens, at the chalets (Hatos) : potatoes and
barley, the latter usually only as high as 6300' ; at a single spot on the
southern slope as far as 7600'. There are no fixed dwellings; the land
is only used for pasturage, without, however, affording the same nourishment
to cattle as other mountains, for connected portions of meadow-turf are rare,
and even here shrubs and thorny plants cover the greater portion of the
slope. Formations of the Central European region : a. Shrubs of Sarothamnus
scoparius, Genista ramosissima, and Quercus Toza, ascending to 6000' ; at the
chalets, these are replaced by thickets of Rosa canina and Berberis vul-

BOTANICAL GEOGRAPHY. 449

garis. b. Thin forests of Pinus sylvestris (5000' 6500'), of small extent on
the Sierra Nevada, the trees being only from 20' to 30' in height ; on the
Serrania de Honda, the Pinsapo forest previously mentioned with isolated
trees of Taxus (5000' 6000'). The Sierra de las Almijarras, south of the city
of Granada, is also partly covered with fir trees up to the summit (vid. sup.),
hence the present forests appear to be only the remains of the zone of Coni-
ferse, which once covered the whole of these mountains, and is now destroyed.
In the nuviatile valleys of the Sierra Nevada, isolated groups of trees, forming
the remains of large forests, among which are the following, part of which
only occur as single trees : Sorbus Aria (5000' 6500'), Cotoneaster grana-
tensis (5000' 6000'), Adenocarpm decorticans (4500' 5500'), Acer opulifo-
lium (5000' 6000'), Fraxinus excelsior (3000' 5000'), Salix Caprea (600(7
6500'), and Lonicera arborea (6000' 7000'), which are the tallest trees grow-
ing there. c. Thorny, low shrubs of a stunted-growth form, an isohypsilous
formation with a, which is principally found on a calcareous soil : Erinacea
hispanica, Genista horrida, Astragalus creticus, Vella spinosa, and Ptilotri-
chum spinosum. This region also contains numerous rock plants, particularly
those belonging to the limestone ; lastly, boggy springs, with limited mea-
dows, exist in the valleys, and these are the localities where most of the
Central European species exist. Formations which in my opinion should be
enumerated as belonging to the alpine region : a. Formation of the Piorno
(Genista aspalathoides} . This shrub, which is sometimes locally replaced by
Juniperus nana and Sabina, forms a broad, connected zone of vegetation
( 8000'), and is distributed downwards, like the Rhododendrons, over a tract
which extends for a considerable distance into the forests ( 5500'). b. Isolated
pastures of rigid Grasses occur on the sloping ground between the Piorno-
thickets. They consist of Avena filifolia, Festuca granatensis, and durius-
cula, and Agrostis nevadensis, Among the endemic forms of Boissier's third
region, in addition to those already mentioned, we have the following, e. g.
Sarcocapnos crassifolia, Silene Boryi, tejedensis, and nevadensis, Arenaria
pungens and armeriastrum, Erodium trichomanefol'mm, and 3 other species,
Anthyllis tejedensis and Ramburei, Astragalus nevadensis, Prunus Ramburei,
Saxifraga Haenseleri, Reuteriana, arundana, biternata, and spathulata, Reutera
gracilis and procumbens, Butinia bunioides, Scabiosa pulsatilloides, Pyretkrum
radicans and arundanum, Senecio Boissieri and elodes, Haenselera granatensis,
Odontites granatensis, Thymm granatensis and membranaceus, Teucrium fra-
gile, compactum, and other species, and Passerina elliptica and nitida.

Fourth region (region of snow), 8000' 11,000'. Isolated patches of snow
never entirely disappear : a connected layer of snow covers the ground for
at least eight months. The soil is constantly kept moist, even in the summer,
by the melting snow. Chalets are no longer met with, although cattle are
driven up to this elevation. The vegetation consists of alpine herbs and
grasses. Only four species of low shrubs belong decidedly to this region,

450 BOTANICAL GEOGRAPHY.

but of these Ptilotrichum spinosum and Salix hastata are extremely rare ;
the two others, Faccinium uliginosum and Reseda complicata do not raise
their woody stem from the ground. The alpine meadows are called " Bor-
reguiles," and they form here a fine sward of Nardus stricta, Agrostis
nevadensis, Festuca Halleri, and duriuscula : and upon this turf, Leontodon,
Ranunculi, Gentians, and other alpine plants grow. In other cases, herba-
ceous plants, growing in the form of tufts, preponderate, and displace the
grass-plat ; as Silene rupestris, Arenaria tetraquetra, Potentitta nevadensis,
Artemisia granatensis, and Plantago nivalis. The alpine rivulets arise from
small lakes, as also from a single spot of glacier-ice, near which, in moist
defiles, we meet with taller herbaceous plants, as Eryngium glaciale, Carduus
carlinoides, and Digitalis purpurea. Lastly, come the plants of the loose
drift, as Papaver pyrenaicum, Ptilotrichum purpureum, Viola nevadensis, &c.;
as also of neighbouring rocks, e. g. Androsace imbricata, Draba hispanica,
Arabis Boryi, and Saxifraga mixta. The following plants are endemic, in
addition to those already mentioned: Ranunculus acetosellifolms and demissus,
Lepidium stylatum, Silene Boryi, Arenaria pungens, Bunium nivale, Meum
nevadense, Erigeron frigidus, Leontodon Boryi and microcephalus, Crepis
oporinoides, Jasione amethystina, Gentiana Boryi, Echium flavum, Linaria
glacialis, Holcus ctespitosus, Trisetum glaciale, Festuca pseudoeskia and Cle-
mentei. Boissier gives the following proportions of the flora of Granada,
which are valuable in a statistical point of view. The following are the
families containing most species in the systematic section of Boissier's work :
239 Synantheracea3 (containing 80 Cynaraceae, 65 Cichoracese, 64 Senecio-
nidese, 29 Asteroidese, 1 of the Eupatorinese), 202 Leguminosse, 164 Gra-
minacea3, 105 Criiciferee, 97 Umbelliferse, 95 Labiatse, 90 Caryophyllacese
(comprising 39 Silenacese, 31 Alsinacese, and 20 Paronychiese), 63 Scrophula-
riaceae, 38 Cistinese, 38 Ranunculaceas, 37 Rubiacese, 36 Boraginacese, 34
Chenopodiacese, 33 Rosacese, 33 Liliacese, 32 Cyperaceee, and 30 Orchidacea3.
Statistical sketch of the four regions assumed by Boissier. In the first
region, 1070 species were observed, of which one sixth only occurred also in
the second region, and a few plants only in sunny places in the upper region.
Of this total number, 542 species are 0, 442 l and 46 Q ; as far as I am
acquainted, this is the only region at present known, in which the number
of annuals is as large or larger than that of the perennials. Of the 442 % ,
19 are trees (vid. sup.), 58 are shrubs less, and 68 more than 3' in height,
the remainder are herbaceous plants. Of the shrubs, 22 are Leguminosse (14
GenistesB), 14 Cistinese, 13 Labiatse (low under-shrubs), 6 Chenopodiacea3,
4 Asparagese (2 Smilax), 4 Amentacese, 4 Solanese, &c. The region con-
tains 860 Dicotyledons, 200 Monocotyledons, 10 vascular Cryptogamia,
distributed through 82 families, of which the following contain most species :
Leguminosse (147), Synantheracese (124), Graminaceae (106), Cruciferse (47),
Umbellifero3 (47), Labiatse (46), Caryophyllaceae (46), Chenopodiaceee (33),

BOTANICAL GEOGRAPHY. 451

Scrophulariaceae (26), Cistineae (21), and Boraginese (20). Cryptogam ic
plants are very rare, because the dryness does not suit the Mosses, nor the
limestone the Lichens. According to its geographical distribution, the flora
of the first region resolves itself into the constituents : a. About 200 are
endemic to Spain, or consist of species distributed as far as Barbary or
Provence ; 12 species only are also found in the East. Characteristic families :
12 Cruciferse (half of which are Brassicacese), 20 LeguminosEe (13 of which
are Genistese)* 24 Synantheracese (11 consisting of Cynaracese), 12 Scro-
phulariaceae (8 Linarite], and 13 Labiatse. b. About 770 Mediterranean
plants, the distribution of which is more extensive, but confined to the
Mediterranean Sea. c. 200 Central European plants, most of which are
either ruderal or marsh plants. Boissier observed 698 species in the second
region, of which one seventh ascend into the third region, and some still higher.
They consist of 202 0, 465 %, and 31 0. To the 1 belong 21 trees,
43 tall and 68 low shrubs : of the tall shrubs, 11 consist of Leguminossc
(10 Genistese), 4 Cistinese, 4 Caprifoliaceae, and 4 Rosacese ; of the under-
shrubs of the Tomillares, 13 consist of Labiatae (4 species of Tkymus), 12
Synantheracese (5 species of Santolina), 7 Cistinese, 7 Leguminosae (Genistese
and Astragalus tumidus), 4 Ericaceae, 4 Chenopodiaceae, and 3 Thymeleaceae.
The region contains 597 Dicotyledons, 93 Monocotyledons, and 8 Ferns, dis-
tributed through 65 families, the following of which contain most species :
Synantheracese (97), Leguminosae (50), Labiatae (44), Cruciferse (41), Um-
belliferae (40), Graminacese (36), Scrophulariacese (27), and Cistineae (23).
Cryptogamic plants are common, tree-lichens commence at 3300' in the
Serrania de Ronda. The second region contains the following components,
arranged according to their geographic distribution : a. 220 Spanish plants,
of which 22 are distributed as far as Provence; 9 species are also
found in the East. Characteristic families : 15 Cruciferse, 15 Leguminosse
(11 Genistese), 15 Umbelliferae, 33 Synantheraceae (19 Cynaracese), 1 5 Scro-
phulariacese, and 1 7 Labiatse. b. About 220 Mediterranean plants. Boissier
collected 422 species in the third region: of these 333 are Tt, 78 , and
11 . To the 1^, belong 14 trees, 44 mostly low shrubs : among these
9 Labiatae (Thymus 5 species), 8 Leguminosae (6 Genistese, 2 Astragaleae),
5 Rosacese, and 4 Thymeleacese. The region contains 358 Dicotyledons,
54 Monocotyledons, and 10 Ferns, distributed through 52 families, among
which the following contain most species : Synantheracese (55), Legurninosas
(29), Graminacese (29), Cruciferse (29), Caryophyllacese (29), Labiatas (27),
Scrophulariaceae (24), and Umbelliferse (20). Arranged according to their
geographical distribution, the third region contains : a. 182 Spanish plants,
of which 101 species appear at present to be confined to Granada. Cha-
racteristic families : 12 Cruciferae, 14 Caryophyllaceae, 15 Leguminosse, 21
Synantheraceae, 12 Scrophulariacese, and 16 Labiatse. b. 185 Central Euro-
pean plants, c. 55 Mediterranean plants. In the fourth region, Boissier

452 BOTANICAL GEOGRAPHY.

found 117 species, of which one third also occur in the third. Of these,
5 only are 0, 3 Q, and 109 l ; moreover, 97 Dicotyledons, 16 Mono-
cotyledons, and 4 Eerns, distributed through 34 families, of which the
following contain most species: Synantheracese (16), Graminacese (11),
Cruciferse (11), Caryophyllacea3 (8), Scrophulariacese (8), Ranunculaceae (5),
and Gentianacese (5). Of Cryptogamic plants, Lichens growing upon rocks
are common. Arranged according to geographic distribution, the fourtli
region contains : a. 45 Spanish plants, of which 30 species are at present
peculiar to the Sierra Nevada ; 13 species are also endemic to the Pyrenees.
b. 66 alpine plants, part of which also occur in plains of the north of
Europe, c. 6 species, which the Sierra Nevada contains in common with
other mountains of the south of Europe.

Wilkomm's botanical letters on his travels (vid. supra),
relate to the whole of Andalusia, and serve both to confirm
and complete Boissier's systematically developed descrip-
tion.

The German traveller at once remarked, that the Sierra Nevada was much
more bare and poorer in shrubs than the other mountains of Spain. He
found, with Boissier, that the northern slope was richer in plants and more
humid than the Alpujarras. In the Serrania de Konda, he saw the forests
of Pinsapo, a tree which unites the growth of the pine with the bark and
arrangement of the branches of the red fir, but which differs greatly in the
thick and short leaves. At a remote period, a great portion of the Serrania
was covered with Pinsapo forests, but the trees have gradually been so cut
down, that the Pinsapo is only now seen as a tree on elevated spots ; it is,
however, found as a shrub from 3000' downwards. The Sierra Tejada, also
a dolomitic mountain between Granada and Yelez Malaga, was formerly
covered by forests of Taxus, from the Spanish name of which, Tajo, the
mountain derives its name. Now isolated trees only occur there at the
source of the Tagus (Fuente del Tejo). The low eastern prolongation of the
Sierra Tejada, the above-mentioned Sierra de las Almijarras is still partly
wooded, between Motril and Granada, with pine trees and oaks, consisting
of Pinus pinea, halepensis and Pinaster, Quercus Ilex and lusitanica. The
slope of this mountain-chain, towards the coast, is also the native place of
Catha Europcea, a shrub, which is found common between Nerja and Motril.
The mountains on the eastern portion of Granada appear to agree in their
vegetation most with the Sierra Nevada ; as does also the barren Sierra de
Alfacar (7000'), which separates the fertile Vega of Granada from the waste
and arid elevated plain around Guadix: Lavandula spica, with some'Cisti, there
forms the covering of the bare slope. The vegetation of the plains of
Guadix and Baza is that of a haloid and gypseous soil. On the boundary
near Murcia, we next meet with the high limestone mountain of Sagra, at

BOTANICAL GEOGRAPHY. 453

Huescar (almost 8000'), where there exist large numbers of pines (Pinm syl-
vestris) : the vegetation here also appears to resemble that of the Sierra
Nevada, inasmuch as Lonicera arborea, e. g. is found there. The same
applies to the Sierra de Eilabres at Almeria (7000'), where Willkomm met
with a number of plants endemic to the Sierra Nevada.

The low plain of Andalusia, or the district of the valley of the Guadalquivir,
is inferior to the highlands of the south as regards the abundance of plants
it contains, but is at the same time more carefully cultivated, especially
around Seville. Willkomm found the tracts lying waste between Seville
and Huelva, covered with dwarf-palms, also woods of pines and cork ; oaks
were common. In the autumn, numerous Liliaceous plants generally
flowered there, with Carenoa lutea B. (Pancratium hnmile Cav.), one of the
Amaryllidaceae. Along the sandy coast, within the lagunes and salt-marshes,
which at Huelva e. g., extend inwards to a considerable distance, pine forests
extend from the Straits of Gibraltar to the mouth of the Guadiana, the
underwood of which, opposite the coast-branches of the Sierra Morena,
consists of Cistus ladaniferus, Ulex Boivini, &c. On the eastern terminal
point of this line of coast, on the Sierra of Algesiras, the traveller met with
a splendid forest of tall cork-oaks and olive trees, such as does not exist
elsewhere in Spain, where Rhododendron ponticum is also mentioned as
occurring by Boissier.

Willkomm visited Algarve in February 1846. The inhabitants distinguish
three regions in it. a. The sandy line of coast (Cousta), which scarcely
extends two leagues into the country, was originally a desert ravaged by the
sea, but by industry has been converted, especially nearTavira, into a paradise-
like district of gardens, containing plantations of southern fruits, vineyards,
and corn-fields. Between Faro and Albufeira, this cultivated surface is inter-
rupted by an extensive pine forest, containing Erica umbellata. Characteristic
plants : Empetrum album, Ulex Boivini and genistoides, Myagrum iberioides,
Arenaria emarginata, Linaria praecox and Unogrisea, Aristolochia batica
(glauca Brot.) and Scilla odorata and pumila. b. The hilly country (Bar-
rocal), extending to 1000', is very much divided, and consists of various
calcareous conglomerates ; it is also fertile and well watered ; still a con-
siderable extent of the good soil lies waste, and is covered with Montebaxo.
The vegetation was still backward; at Louie, e. g. Erica austmlis
and lusitanica, Osyris quad^artita, and several Narcissi, were found.
c. The mountainous region (Serra), a terminal, undulating prolongation of
the Sierra Morena, like the latter consisting of grauwacke and clay-slate,
the western portion only, the Sierra de Monchique, being composed of granite
and basalt, appeared dark green, yet not susceptible of cultivation. It is a
remarkable fact, and tends to show the great influence of the geological
substratum, that even here the shrubs of the Spanish Sierra Morena pre-
dominate, Cistus ladaniferus being very common, but mixed with the two
Erica of 4he Barrocal. The valleys of the Sierra de Monchique are, however,

454 BOTANICAL GEOGRAPHY.

wooded with chesnut trees and cork-oaks, with which Rhododendron ponticiim
grows in common. This mountain does not ascend more than 3800' accord-
ing to Portuguese measurements, but the upper vegetation is subalpine, and
corresponds with the altitudes of 5000' 6000' of Andalusia, which, however,
I should not ascribe, as the author does, so much to the influence of the
capacity of the granite for heat, as to the cessation of the influence of the
plateau. The depression of the climate below the natural standard is not
occasioned by storms, but the adjacent sea and low country cause a normal
fall of the temperature in a vertical direction ; whence Andalusia and the
whole of Spain are abnormal in this respect, and the limits of vegetation
extend to a disproportionate height.

Schouw has now published his work upon the Coni-
ferous trees of Italy (see Ann. Rep. for 1844), in greater
detail (Ann. Sc. nat. 1845, torn, i, p. 230).

Districts of the distribution of the species : 1 . Pinus sylvestris L. (includ-
ing P. uncinata D. C.) South slope of the Alps, 6000' below 1000' at
Tagliamento, an Apennine of Montserrat. 2. Pinus Pumilio, Hk. South
slope of the Alps, 4000' 7500'. 3. P. magellemis Sch. (P. Pumilio Ten.
and MugTius Guss.) appears to hold the same relation to 4 as 2 to 1 . The
Abruzzi at Majella, 5000' 8300'. 4. Pinus Laricio Poir. (P. sylvestris and
nigrescens Ten.) forms the forest of jEtna, 4' 6000'. Calabria and Abruzzi
at Majella. Pinus nigricans Host, and P. Pallasiana are probably the
same, as I have also assumed. 5. Pinus Pinaster Ait. Apennines. 0' 2800'
on Monte Pisano. 6. Pinus Pinea L. Apennines as far as Genoa, with
5. 0' 1500' in the Western Apennines, 2000' in the south of Italy. 7. Pinus
halepensis Lamb. The whole of Italy, as far as the Apennines ; 0' 2000'
on the Somma at Spoleto. 8. P. brutia Ten. Calabria, on the Aspromonte
at Reggio, 2400' 3600'. It does not, however, appear to me sufficiently
distinct from Pinus Laricio. 9. Pinus Cembra L. Alps, 4 6500'. 10. Abies,
excelsa D. C. (P. Abies L.), Alps ; 1000' (Tolmezzo) 7000' (Stilfser Joch).
11. Abies pectinata D. C. (Pinus Picea L.) Prom the Alps to Madonia;
1000' 4500' in the Alps, 5500' in the Apennines. 12. Larix europaea.
Alps; 1500' (at Piave) 7000'. 13. Cupressus sempervirens L. Alps to
Sicily, 0' 2500'. 14. Juniperus communis L. South to 40; Alps,
0' 5000'. 15. Juniperus nana. W. Alps, 5' 7500'; Apennines. 16. /.
hemisphcerica Prl. .2Etna, 5' 7000'; Calabria. 17. /. Oxycedrus L.
Apennines, from 1000' 3000'. 18. /. macrocarpa Sibth. Along both the
seas from Pisa to Sicily. 19. /. Sabina L., Alps; Apennines. 20. /. phce-
nicea L. Along both the seas from Nice to Sicily. 21. Taxus baccata L.
Alps; Apennines.

Parlatore has commenced a Mora of Palermo (Flora
Palermitana, vol. i. Firenze, 1845, 8vo). The first volume
contains only the Grasses (130 species), with very full
descriptions.

BOTANICAL GEOGRAPHY. 455

New species : Avena Heldreichii, Melica nebrodensis, Vulpia panormitana
and attenuata (Festuca sicula Mor.)

Regarding the periods of the vegetation of wheat (Tri-
ticum vulgar e hybernum), we find in Damn's work (1. c.
p. 347), the following observations made in the year 1842:

Average period of Sowing. Time of Harvest.
Malta . . Dec. 1 . . . . May 13 . . 164 days
Sicily . . Dec. 1 . (Palermo) May 20 . . 171
Naples . . Nov. 16 . . . . June 2 . . 195
Rome . . Nov. 1 .... July 2 . . 242
Berlin . . 299

Link read some observations upon the vegetation of
Dalmatia, before the Geographical Society of Berlin (Mo-
natsberichte, Bd. 2).

Visiani has had individual plants of Greece and Asia
Minor drawn, from Parolini's collections (Memorie dell'
Institute Veneto, vol. i, 1843) : 6 of theLabiatae (Thymus,
Stachys), 2 Boragineae (AncJmsa, Lycopsis), Dianthm
Webbianus and Sedum Listonia. The new species which
he has there proposed had in part, however, been pre-
viously described elsewhere.

II. ASIA.

Parts 11 18 of the ' Illustrationes plantarum orien-
talium' of Gr. Jaubert and Spach (see Ann. Rep. for
1843) have appeared (Paris, 1844-45). The following
families and genera are described in detail : Polygoneae,
Asarineae, Chenopodeae, Legurninosse, principally Ge-
nisteae (including G. gracilis, t. 143 = G. carinalis, m.),
Cousmia. In Lorent's oriental travels (Wanderungen
im Morgenlande. Mannheim, 1845, 8vo), 35 species are
described by Hochstetter as new, most of them from
Syria and Armenia. Endlicher and Diesing have de-
scribed 6 Algae which were collected by Kotschy in the
Persian Gulf (Bot. Zeit. 1845, p. 268).

Mahlmann has elaborated Chanykoff's observations on

456 BOTANICAL GEOGRAPHY.

the climate of Bokhara (Berliner Monatsber. fur Erd-
kunde, Bd. 2, pp. 132-40).

North winds are always prevalent in Chanat, hence they are in the direc-
tion from the steppe to Hindu-Kusch, which explains its freedom from rain
and its continental distribution of heat. During eight months the wind was
in the opposite direction ten times only. In the city of Bokhara (1116'
above the sea) Chanykoff however found the mean temperature during a
severe winter = 30 F., i. e. lower, with the exception of Pekin, than has
previously been anywhere observed in the same latitude. The trees bud
between the 20th of March and the 10th of April. The vegetation of the
steppes between Samarkand and Karschi lasts from the middle of March to
the end of April ; but the temperature remains high from the middle of
March to the end of November, and is excessive in the summer.

TschihatchefFs work upon his travels in eastern Altai,
principally the district of the course of the Jenisi (Voyage
dans T Altai oriental. Paris, 1845, 4to), contains a list of
the plants collected by the author in a portion of the
district, part of which had not been previously examined ;
when determined by Turczaninow they were found to
agree with those of adjacent countries.

The following were the trees : Larix sibirica, Abies Pichta, Pinns syhes-
tris and Cembra, Alnus viridis, Betula alba, Salix Pontederana, pentandra, and
stipwlaris Turcz., Popuhis alba, tremula, and laurifolia, and Sorbus aucuparia.
The following families and genera have been described (Bull. Moscou, 1845),
as forming an addition to Turczaninow's Flora of the Baikal Regions
(see the preceding Ann. Rep.) : 1 Adona, 1 Cornm, 6 Caprifoliaceee, 7
Rubiaceae, 6 Valerianeae, and 2 species of Scabiosa.

G. Reichenbach has described some Orchidaceae in
Goring's collection from Japan (Bot. Zeit. 1845, p. 333).

The following divisions of R. Wight's illustrated work
upon the flora of Hindostan (Ann. Rep. for 1840) have
appeared according to the advertisement : Vol. ii, part 1
of the Illustrations of Indian botany, with 39 plates
(Madras, 184]); of the Icones plantarum Indiae orien-
talis, the conclusion of the first volume, consisting of 16
parts and 318 plates; vol. ii, with 318 plates (ib. 1840-
42) ; and vol. iii, parts 1-3, with 409 plates (ib. 1843-46).
Wight has also published a Spicilegium IN eilgherense, with
50 plates (ib. 1846, 4to), in which particular plants of
Nielgherry are figured : the latter appears, however, to

BOTANICAL GEOGRAPHY. 457

be only an abridgment of his previous work (see Gardner's
remarks in the Lond. Journ. of Bot. 1845, p. 565). As
stated in a letter, a memoir by Madden upon the Coniferae
of India is contained in the ' Quarterly Med. and Lit.
Journal/ 1845, pp. 34-118, published at Delhi. Gardner,
the Brazilian traveller, who is now superintendent of the
garden at Columbo in Ceylon, has reported upon his
botanical excursions in Nielgherry (1. c. pp. 393-409 and
551-67) ; he enumerates the localities of the plants exist-
ing there.

De Vriese has commenced publishing an illustrated
work upon select plants of the Dutch East Indian posses-
sions (Nouvelles Recherches sur la Flore des Possessions
Neeiian daises aux Indes Orientales. Fasc. 1, with 3
plates. Amsterdam, 1845, fol.) : it contains a description
of some new Styraceae from Sumatra and Java, a figure
of Casuarina sumatrana, as also of the new Pinus Mer-
kmii from Sumatra. Hasskarl has continued his remarks
upon various points relating to the plants of Java both
in the ' Ratisbon Flora' (1845, pp. 225 et seq., containing
the Rubiaceae), as also in V. d. Hoeven's Zeitschrift
(Bd. 12, pp. 77 et seq., comprising the Malvaceae and the
allied families) . Montagne is describing the Lichens and
Mosses of the Philippine Isles, from Gumming 5 s collec-
tions (Lond. Journ. of Bot. 1845, pp. 3-11).

III. AFRICA.

Fresenius has published Contributions to the Flora of
Abyssinia froniRiippelPs collections (Mus. Senckenbergian.
vol. iii, 1845) : containing copious descriptions of those
Polygonese which have already been made known, and
some new Synantheraceae.

He gives, at the same time, a figure of the Lobeliaceous tree of Abyssinia,
Gibarra (Rhynchopetalum montanum = Jibera of the preceding Ann. Rep.),
and has represented its habit as follows : from 6'-7' in height, stem hollow, a
crown of lanceolate leaves and tall bunches of flowers ; hence the dimensions
of the plants observed by Ruppell in Simen, between 11,000' and 12,000'

458 BOTANICAL GEOGRAPHY.

were less in those found by Harris in Skoa. (See the preceding Ann. Hep.
p. 382.) C. H. Schultz has described some of Riippell's new Cichoraceae
from Abyssinia (loc. cit. p. 47).

Endlicher and Diesing are describing new Algae from
the Natal Colony (Bot, Zeit., 1845, pp. 288-90).

IV. AMERICA.

Seller has made some isolated systematic remarks upon
a collection of plants from the coasts of Davis's Straits
and Baffin's Bay, in the ' Annals of Natural History '
(vol. xvi, pp. 166-74).

Forry has compared the results obtained at the meteoro-
logical stations of the United States since 1819, and
traced the distribution of heat in various points of view
(Amer. Journal of Science, 1844, extracted into the
Biblioth. de Geneve, vol. Ivii, pp. 140-50).

The unusual, nay, unparalleled accumulation of fresh water in the Canadian
lakes, which, at a mean level of 1000', include a surface of almost 4000
square geog. miles, procures for the northern states an insular climate for a
very considerable distance into the eastern forest-region. Hence the difference
between the summer and winter does not become excessive until we
arrive just beyond the Mississipi, as also between the lakes and the
Atlantic Ocean ; in Lower Canada, e. g. the extremes of temperature are
somewhat greater than in Michigan on the one hand, and the coast of Nova
Scotia on the other. In the southern states, the annual curve resulting from
the influence of the two oceans becomes still less arched than in the north,
until, in Florida, it gives way to an almost tropical uniformity. The diffe-
rence between the temperature of the summer and winter amounts there at
Key West to only 43 F. ; flowers bud there throughout the year without any
general winter-sleep . During a space of six years th e thermometer never rose,
at this station, above 89'6F., and never sunk below 44' 6 F. The atmo-
spheric precipitations are unequally distributed in Florida : in the central
districts there are 309 fine days in the year, on the coast 250, and at the
lakes, in the northern part of the state, only 11 7; but the air generally
abounds in moisture, and the formation of dew is common.

Macnab has continued the Botanical Report of his
travels (Ann. Nat. Hist.,, xv, pp. 65 and 351). Berkeley
has published an account of some new Fungi from Ohio
(Lond. Journ. of Bot., 1845, pp. 298-313).

BOTANICAL GEOGRAPHY. 459

Geyer's reports upon the characters of the vegetation
of the prairies on this side of and beyond the Rocky
Mountains (Lond. Journ. of Bot., 1845, pp. 479-92, and
653-62), in connexion with Fremont's investigations
(vid. inf.), are immediately connected with the descrip-
tions given by the Prince of Wied, to which the former
are far inferior in regard to their too aphoristic style,
but are as superior in systematic botanical knowledge.

The traveller ascended the Platte River from the State of Missouri,
through the Osage district, as far as its source in the Rocky Mountains,
traversed the mountains and the Colorado of California at about the forty-
second parallel, and thus arrived at the Oregon territory. The western
and southern limit of the prairies, which, to the south of Arkansas (accord-
ing to De Mofras' map), are connected with the forests of New Mexico
(37) are not far from the Lower Kanza, in the district of Osages
(39 N. lat.) Hence even here the forests of the valleys along the
river become more numerous, the prairies abound to a greater extent in
flowers, and the period of the summer drought is shortened. The most
common species among the deciduous trees of Illinois, which are almost
the same as those mentioned by the Prince of Wied (Ann. Rep. for
1842), and which also form the forests of the banks of the river on the
Lower Missouri, gradually meet here with their western boundary, and they
diminish in height the nearer they approach the sandy valley of the Platte.
The herbaceous plants of this fertile prairie, however, become proportion-
ately more numerous, and produce an uninterrupted succession of flowers
throughout the spring and entire summer. In April isolated spring plants
appear ; in May and June the whole of the undulating surface for an immense
distance is in flower, the plants consisting, e. g. of Amorpha canescens,
Batschia, Castilleja, Pentstemon, Cypripedium candidum, &c. ; taller herbaceous
plants follow: Petalostemon, Baptisia, Phlox aristata, Asclepias tuberosa,
Lilium canadense, and Melanthium mrginicum ; and finally, in the latter part
of the summer, almost exclusively Synantheraceae, from tall Helianthese
down to the dwarf Aster sericem.

The rich soil of the prairies terminates at the river Platte, with the lime-
stone of the Missouri, which favours the vegetation detailed above. The
lower terrace follows next ; it is 900'-1000' in height, and further up the
stream is connected with the elevated surface of the upper steppe. The
stony and sandy crust of the earth is formed of the detritus of granite, which
is expanded over horizontally laminated sandstone and bituminous slate. The
woods on the islands of the river then consist of Popuhis canadensis, Ulmus
americana and/w/m, Negundo and Celtis occidentalis ; on the bank, thickets
of Salix long i 'folia, with Amorpha frutescens, Rosa parvi folia, Rubus occiden-

460 BOTANICAL GEOGRAPHY.

talis, and Rhus glabriim, predominate on the open prairie, which in May and
June is rendered moist by the atmospheric precipitations : the vegetation,
nevertheless, scarcely lasts longer than these short weeks of spring. The
following forms may be mentioned as characteristic of the flora of the prairie;
they are subdivided according to their localities, although the author has not
arranged them in the form of a summary : Of the Leguminosae, Astragalus,
e. g. A. adsurgens and caryocarpm, Oxytropis, Phaca, Petalostemon, Psoralea,
Glycyrrhiza, and Schrankia', Malvaceae, Sida coccinea ; Cactacese, Mamil-
laria simplex and Opuntia missurica ; Onagrarise, (Enothera and Gaura ;
Synantheracese, principally Helianthese, e. g. Echinacea, Rudbeckia, Heliopsis;
moreover, Artemisia, e. g. A. caudata, and Lygodesmia ; Scrophulariacea3,
Pentstemon and Castilleja; Hydrophyllacese, Ellisia ; Boragineae, Batschia;
Nyctaginese, Calymenia ; Liliacese, Yucca ; Graminaceae, e. g. Sesleria dac-
tyloides, Crypsis, Stipa, Agrostis, Eriocoma, &c.

The remaining large district is denominated by Geyer the upper saline
desert region, the area of which extends far inwards symmetrically, on both
sides of the Rocky Mountains, from Missouri to Lower Oregon, a desert
elevated surface resting upon sandy rocks, and gradually ascending from
1200' to more than 4000'; so that the chains of the Rocky Mountains, in
spite of their elevated central ridges, cannot by any means be regarded as
forming a boundary of vegetation. The boundaries of this immense steppe,
which everywhere affords pasture, Geyer considers as formed, in the north, by
the Saskatchawan and Lake Winnipeg ; in the east (the same as the Prince
of Wied), by a line running longitudinally through loway, or the former dis-
trict of the Sioux (Great Sioux river and Moine's river) ; in the south, by the
Upper Arkansas ; in the west, by the mouth of the Wallawalla, in the Oregon
district (more distinctly by Eremont, the union of the two principal forks of
this river, the Lewis river and the Upper Columbia) ; hence about 3S-54 N.
lat. and 77-101 W. long, from Ferro. With the exception of the pine
and snow -clad central chain of the Rocky Mountains, this space contains no
forests. The prevailing character of the flora is generally the same as that
described by the Prince of Wied, that of the Upper Missouri. Beyond
the Rocky Mountains also, as in the district of the source of the River
Platte, the steppe is covered with two social shrubby Artemisias (Art. triden-
tata and cana). The Pulpy -thorn, Sarcobatus vermicularis (S. Maximiliani
N.), also called the Salt-cedar, is found everywhere on the saline soil as low
down as Oregon ; it is a shrub, with numerous stems from 3'-8' in height,
with diverging thorny branches and dark-green succulent leaves. Considering
the similarity of the climate and soil of the prairies and the Russian steppes,
it is an interesting fact, that this genus, which was first recognised as distinct,
from the examination of Wied's collections, according to both Lindley and
Torrey (Fremontia ej., Satis vermicularis, Hook.), is a true member of the
Chenopodiacea3 (Lond. Journ. of Bot., 1845, pp. 1 and 481), and grows in

BOTANICAL GEOGRAPHY. 461

common with other Halophytes belonging to the same botanical group. The
other most common thickets of the upper steppe consist of Maeagnns
argentea and Shepherdia argentea ; then Amorpha frutescens, Rosa parvifolia,
and woody Synantheracese, e. g. Iva and Bigelovia. Juniperus andina (J. repens
of Wied), with Yucca angustifolia, appear to be confined to the Missouri
country below the mouth of the Yellowstone river. Geyer's further distinc-
tions of several districts of vegetation in the region of the upper terraces
has not been carried out sufficiently clearly. The following may be regarded
as characteristic forms : Of the Leguminosse, Astragalus, Homolobus,
Psoralea, Glycyrrhiza, Hosackia, Schrankia, and Amorpha; Cruciferee, Stan-
ley a pinnatifida ; Loasese, Bartonia ornata ; Onagraria?, (Enothera ; Cacteee,
Opuntia missurica ; Umbelliferse, Cymopterns ; Synantheraceae, in addition
to the above-mentioned shrubs, several Chrysopsidese, Cichoracese, Achillea ;
Scrophulariaceae, the same genera as those of the lower terraces ; Chenopo-
diacece, in addition to Sarcobatus: KocMa, Salsola, Chenopodium, and Atriplex;
Liliaceee, CalocJiortus and Allinm, Iris, Triglochin maritimum, Carex ; Gra-
minacese, e. g. Triticum missuricum, B ordeum jubatum, and Ceratochloa.

Geyer's description is rendered geographically more clear by the excellent
diary kept by FREMONT of his travels, who, being the chief of an expedition
of discovery, and furnished with botanical knowledge, explored the whole of
the steppes of the North American prairies, down to Lower Oregon and
Upper California, in different directions, with the most fortunate results
(Narrative of the Exploring Expedition to the Rocky Mountains in 1842, and
to Oregon and North California in 1843-44. Washington, 1845. I am only
acquainted with it from the English edition, London, 1846-8). On this side
of the Rocky Mountains, Eremont followed the same course to the River
Platte as Geyer ; on the second occasion, he ascended the Kanza and its
accessory streams, to the central chain. The country ascends very gradually
from the bifurcation of the Kanza (79 W. long.), to the foot of the Rocky
Mountains, and on the west side of the mountain the land sinks to the con-
flux of the Lewis and Oregon, as is evident from the following line of level
which was determined barometrically by Eremont, and intersects the entire
steppe from east to west. Bifurcation of the Kanza (79 W. long.) = 926';
3iver Platte (81) = 200(y; River Platte (83) ==2700'; Eort Laramie, on
the Platte (87) = 4470'; and almost in the same meridian, Eort Trains
(40 16' N. lat.) == 4930'; as also the River Arkansas (38 15' N. lat.)
= 4880'; Artemisia-steppe, at the eastern foot of the Rocky Mountains
(41 36' N. lat. and 90 W. long.) = 6820'; south pass through the Rocky
Mountains, in a deep depression, which does not possess any mountain cha-
racter (42 27') = 749CK; foot of the Rocky Mountains, at the upper part
of the course of the Colorado of California (41 46') = 6230'; Eort Hall,
on the Lewis (43 N. lat., 95 W. long.) = 4500'; River Lewis (43 49'
and 99) 2100'; River Lewis (44 17' and 100 W. long.) = 1880'.

462 BOTANICAL GEOGRAPHY.

The open prairie-steppe beyond the Rocky Mountains is covered generally
with shrubs of Artemisia, between which, however, cattle everywhere find
food in nutritive grasses. Purshia tridentata, one of the Spiraeacese, which
frequently accompanies the Artemisia, is a shrub peculiar to this part. The
nutritive plants used there by the Indian hunters in cases of necessity, cor-
responding to the Psoralea esculenta on the Missouri, consist of Valeriana
edulis (Tobacco-root), Cirsium mrginianum, a species of Anethum (Yampeh),
and Kamassa (Kamas), Fr. indescr. The bank-forests of the cotton-wood
(Populus) are not met with until we arrive at the lower regions : they appear
to be entirely wanting on the upper terrace. At the bifurcation of the
Oregon, where the prairie terminates (101 W. long.), the wooded promon-
tories of the western chain of high mountains commence, which may be
compared to the Rocky Mountains in extent, and projecting everywhere
above the snow-limit, probably exceed them in height. Being a continua-
tion of the Californian Andes, it is called, in Upper California, Sierra Nevada,
in Oregon, the Blue Mountains, and the Cascade-chain, where, on the south
side of the united rivers, near Fort Vancouver, it rises into high snow-moun-
tains, as at Mount Hood. At Oregon, the forests of this high moun-
tain-chain (explored between 2700' and 3800'), which are only interrupted
by the most splendid meadow-slopes, consist of birch, but above all of
various Coniferous trees remarkable for their enormous dimensions, which
are such as are not met with in any other part of the globe. The larches
were sometimes 200' high (p. 182), the firs were of the same height, with
stems T in diameter ; in the former, the unbranched stem beneath the crown
was sometimes 100' in length. "White spruces which gave off branches
down to the root appeared nevertheless to measure 180', perhaps 200'.
The Cascade-chain separates the mild climate of the western coast of the
Oregon district from the dry prairies equally as definitely, but in an inverse
direction, as the Peruvian Andes do the west desert coast-region from the
more humid higldands. This meridional mountain-chain, which intersects
the River Columbia about twenty-five or thirty miles from its mouth, receives
the mists and rain which are driven over to it from the Pacific Ocean, but
which do not penetrate to the clear sky of the steppe. At the rapids of the
Columbia, the " Dalles" within the mountain-line, the rainy season is already
unknown, which on the coast denotes the winter, and this season is only
recognisable there (45 N. lat.) by a slight fall of snow, which scarcely lasts
on the ground for two months. The cause of the winter rainy season at the
mouth of the Oregon, where west winds predominate, appears to me to
depend simply upon the fact, that in the summer the sea, whilst in the winter
the continent, is the coldest, so that during the latter period of the year, the
humid winds from the sea must quickly lose their moisture in passing over
the coast-district. But the steppe lying behind the mountains is a highland ;
as such it exceeds the coast in warmth and dryness, and cannot, therefore,

BOTANICAL GEOGRAPHY. 463

readily precipitate the aqueous vapour from the westerly current of air.
The same, however, holds good here, from whatever other points of the
compass the wind may blow, so that instead of steppes, deserts would be
expanded between the Rocky Mountains and the California!! Andes, if this
internal country were not also so copiously watered by these mountains, and
thereby also subject to local precipitations. The climatal relations of the
Oregon-district also perfectly explain the drought of the prairies in the Mis-
souri described by the Prince of Wied.

Prom Columbia, Fremont went to the eastern foot of the Sierra Nevada,
as far as the 39th degree of south latitude, following the boundary-line be-
tween the steppe- and the forest-regions. Below the 42d degree, at the
south water-boundary of the district of the Oregon-river region, the inland
country is elevated into a mountain-chain running from east to west, and not
void of forests, and this appears to form a connexion between the Californian
Andes (S. Nevada) and the Rocky Mountains.

South of the chain, a desert highland is situated ; it is probably for the
most part uninhabitable, and, from the nature of its soil and its declivity, it may
be compared with the uninhabitable regions of Persia ; it ought to be called
the Californian salt-desert (Fremont's great interior basin). An Indian guide
pointed to it, saying at the same time, " There are the great llanos no hay
agua, no hay zacata, nada" i. e. Plains without water, without herbage :
" Every animal that enters it must die." Entirely surrounded by mountains
which form its borders, bounded to the north by the river-limit of the
Oregon, to the south by a similar chain, covered with snow, towards the
Colorado, and on both sides by the Sierra Nevada and the Rocky Mountains,
it only contains internal streams, which lose themselves in the desert or
in salt-water lakes, and is perhaps dry and destitute of springs for the space
of many days' journey. As the greater part has not hitherto been explored
by any traveller, we are confined, with regard to the altitudes, to the follow-
ing measurements made by Fremont, which, in fact, only relate to the
external margin : on the plateau of the great salt-water lake Utah (41 30' N.
lat. and 95 W. long.) = 4200'; Lake Pyramid, at the foot of the Sierra
Nevada (39 51') = 4890'; foot of the Sierra Nevada (38 50') = 5020';
on the boundary-mountains, Bear River, on the slope of the Rocky Mountains
(42 and 93) = 6400' ; pass from the Bear River to Colorado (41 39') =
8230'; pass over the ^Sierra Nevada to the Bay of St. Francisco (38 44')
= 9338'. The salt-desert differs from the prairies of Missouri, as from the
Artemisia-steppes of Oregon, by its excessive dryness, rocky soil with vol-
canic heaps, by the more general presence of salt in the soil, and, as a
consequence of these conditions, by the absence of the growth of nutritive
grasses ; nevertheless, from the strength and number of the rivers which
enter it from the boundary-mountains, we may conclude as to the existence
of oases on its streams. The vegetation consists almost exclusively of

464 BOTANICAL GEOGRAPHY..

shrubby Chenopodiacese, with which in tracts Artemisias are mixed, and along
the Sierra Nevada, and to the south of the 41st degree of latitude, Ephedra
occidentalis, forming an evergreen shrub. The most common of the Cheno-
podiacese found here is also Sarcobatus vermicularis ; Obione is next mentioned,
of which Obione rigida Torr. and Fr., with another new species, occurred at
Utah ; Salicornia also covered the banks of this lake. The woods of the
boundary-mountains to the north of Utah consisted of deciduous trees :
Populus, Salix, Quercus, Cratagus, Alnus, and Cerasus. Below the 39th de-
gree of longitude, the Sierra Nevada was crossed with great difficulties in
the depth of the winter, so as to reach the valley of Sacramento. The lowest
forest-zone on the desert side of the mountains consisted of a pine, the seeds
of which were edible, Pinus monophylla Torr., a tree from 12' to 20' in height,
and with a stem at the most 8" in diameter, which, with some roots and the
salmon found in the waters, form the food of the Indians. Further upwards
this pine (nut-pine) was found somewhat larger, its diameter amounting to
15". But it was not until an altitude of 6000' had been reached, that
Coniferous forests of a taller growth and of a different species were met with,
accompanied by a more luxuriant vegetation, in which the first indications of
a fairer climate were met with, At 8000' the trees were almost as gigantic
as in Oregon : red pines as high as 140' and 10' in diameter (Pinus Colorado
of the Mexicans) predominating, and with them tall cedars 130' in height,
and two species of fir of an equally tall growth (white spruce and hemlock
spruce). Trap-rocks form the fertile soil of these splendid tall forests, to a
considerable depth. On the west side of the mountains, below the Coniferous
zone, Fremont arrived at a region of evergreen and other oaks, which corre-
sponds with Hinds's representation of the character of the region of St. Fran-
cisco : here, after the impressions left by the deserts, the traveller was de-
lighted with the most luxuriant spring flowers in the valleys of the Sacramento
and St. Joachim.

On his return, Fremont crossed over the Californian Andes by a much
lower pass, below the 36th degree, and travelling parallel with the Colorado,
on the southern border of the salt-desert, returned to the Great Salt-lake and
the Rocky Mountains. This road, which forms the course taken by the
caravans in going from New Mexico to California, was rocky and mountainous
(sloping off towards the Colorado from about 5000' to 2000') : the vegetation
was scanty, corresponding to the character of the flora of California. A
tall Zygophyllaceous shrub (T^ygoph. calif or nicum Torr., Fr.), a Yucca, and
numerous Cacti constitute the principal forms of plants over extensive tracts ;
and from the north towards this part, as far as the woods of the Yucca, the
Artemisia tridentata of the steppes extends : the traveller would not, however,
give the preference to the former, since the stiff and unsymmetrical form of
the Yucca appeared to him the most repugnant formation in nature. Among
the shrubs of this region, he mentions Ephedra occidentalis, Garrya elliptica,

BOTANICAL GEOGRAPHY. 465

which forms dense thickets on the banks of the rivers, and
odoratum Torr., one of the Mimoseee, 20' in height. The northern forms
distributed thus far were (36 N. lat.) : Pinus monophylla, Purshia tridentata,
and Populus and Salix on the banks of the rivers.

The snow-line of the Rocky Mountains was estimated on the Snow-Peak
(42 43 N. lat.) at 11,800' (i. e. 1800' above the measured point 10,000'),
This mountain, the summit of which, 13,570' high, Fremont ascended, belongs
to the accessory chain of the Wind River mountains, but is regarded as the
most elevated of the entire system. Above the Coniferous region, the alti-
tudinal limits of which were not determined there, it contains a copious
alpine vegetation, which, according to the examples brought forward, are
principally characterised by Hudsonian forms, just as those of the Alps are
by arctic forms.

Fremont's observations upon the tree-limits of the continent of North
America are extremely remarkable ; they show that they are much higher than
in corresponding latitudes of Europe. Not only in the Californian Andes
were the pine forests found to extend above 8000', but on the east side of the
Rocky Mountains, in the district of the so-called Park, in the region of the
source of the southern bifurcation of the River Platte and the Arkansas
(39 20' N. lat.), Fremont found that even at an elevation of 10,430', " the
pine forest continued, and grass was good. We continued our road, occa-
sionally through open pines 3 with a very gradual ascent ; and having ascended
perhaps 800 feet, we reached the summit of the dividing ridge, which would
thus have an estimated height of 11,200' " (p. 314.) Hence the altitude of
the tree-limit of the Rocky Mountains in the latitude of Valencia may be
assumed as 11,000' ; the most elevated tree-limits of the south of Europe,
the isotherms of which are of such very different temperatures, scarcely ascend
beyond 7000'. If the influence of the highlands of North America is so
great in moderating the vertical diminution of the temperature of the summer,
we are justified in anticipating similar phenomena in central Asia. There is
especially, one observation which corresponds with this supposition, and it is
the only one with which I am acquainted, viz. that relating to the valley of
Spiti, in Lesser Thibet, where, according to Jacquemont, at the same altitude,
but a more southern latitude (32 N. lat.), dwarf trees alone occur. But it
is not only the heat which causes the dense tall forests to ascend to such
considerable elevations in North America ; the humidity of the air or of the
soil must also be taken into account. In the south of Europe, the tree-limit
does not ascend in proportion to the increase of heat, since it is frequently
situated at a greater altitude on the south side of the Alps than at any
more southern point of the continent. In Thibet, where the highlands even
ascend to the level of the tree -limit, the limitation of the growth of trees is
not caused by cold, but by dryness. Now, it is a circumstance common
to both the mountain-chains of North America, that under the latitudes

466 BOTANICAL GEOGRAPHY.

of the south of Europe, they ascend far above the limit of perpetual snow.
Here the drying influence of the plateau, which lies far below the forests, is
removed by the masses of snow; not so, however, in Thibet, where the
country corresponding to the plateau ascends to the snow-line. On the
mountains of North America, as on the south side of the Alps, sufficient
water is thawed in summer from the large fields of snow to irrigate the
elevated forests : thus they are provided with a permanent source of moisture ;
even when the prairies are without rain for months, they never dry up,
whilst upon Pindus and the Apennines the winter snow soon disappears ;
whilst in Thibet the melted snow upon the elevated surface evaporates again,
without fertilizing the soil.

In a botanical appendix to Duflot de Mofras' work
upon the western coast of North America (Exploration
du Territoire de 1'Oregon, &c., 2 vols. 8vo. Paris, 1844),
a list of about 300 Calif ornian plants is given. It is,
however, derived from old sources, and is disfigured by
errors of the press to such an extent as to be rendered
almost useless.

In the work itself we find the following statements regarding the course of
the seasons in California : 1. In Upper California, e. g. in the latitude of
St. Francisco (38 N.), the rainy season, with prevailing south-east winds,
lasts from October to March. From April to September north-west winds
blow ; it then never rains, although fogs are common on the coast ; the soil
also then loses its verdure (ii, p. 46). On account of the length of the
drought, the total amount of the atmospheric precipitations is less than in
the south of Europe. 2. The vegetating season of the arid western coast of
Lower California, however (30 to 23 N. lat.), occurs with its atmospheric
precipitations during the summer (i, p. 239). 3. On the eastern coast of
this peninsula, at Cape Lucas, in the Gulf of California (mer vermeille), and
on the north-west coast of Mexico we find an inversion of the trade-wind,
(inversion d'alize, i, p. 171), as south-westerly or westerly winds predominate
there. In Mazatlan (23 12') the rainy season coincides with south-westerly
and westerly, and the dry season with north-westerly winds (i, p. 172) ; the
same is the case at St. Lucas, where these latter monsoons prevail from No-
vember to May (i, p. 229). Within the gulf, where, although it is beyond
the tropic, the monsoons are the same, the amount of rain appears to diminish
very considerably. We cannot imagine anything more miserable and
neglected than these two coasts, which lie waste from a deficiency of water
(i, p. 205).

I find a notice of Plantse Lindheimerianse, by Gray,
which probably contains an account of Lindheimer's

BOTANICAL GEOGRAPHY, 467

collection from Texas; but I am unacquainted further
with the work.

A. Richard and Galeotti intend publishing a monograph
of the Mexican Orchidacese, which will include 460
species ; of these about a third part are new. They have
published preliminary diagnoses of the new species (Ann.
Sc. nat., 1845, t. iii, pp. 15-33). V. Schlechtendal's
contribution to the flora of Mexico, for the present year,
refers to the Asphodeleae (Bot. Zeit., 1845).

Purdie (Ann. Rep. for 1843) has continued his botani-
cal reports from Jamaica (Lond. Journ. of Bot., 1845,
pp. 14-27).

The Cactacese, which are common on the south coast of the island, are
absent from the north side. In the former locality, at Bath, he found coast-
mountains about 3000' high, covered with a tall forest of Podocarpus Pur-
diena Hook., one of the largest forest-trees of Jamaica ; one of them, which
had been cut down, measured more than 100', 40' up to the crown, and at
the height of a man above the root, it was 3|' in diameter. Podocarpus
coriacea occurs above a level of 5000' or 6000'. The coffee-plantations are
situated on the south side of the island, e. g. at the pass from Kingston to
Bath, between 3000' and 6000'. Coffea does not thrive at a greater ele-
vation.

Kuiize has enumerated and described the new species
among the Ferns collected by Moritz in Caracas (Bot.
Zeit., 1845, pp. 281-8). Of Bentham's descriptions of
Schomburgk's plants from Guiana, the Polygonacese
(14 sp.) and Thymeleacese (3 sp.) have appeared, as also
those of the Acanthacese (17 sp.) from Nees v. Esenbeck
(Lond. Journ. of Bot., 1845, pp. 622-37). Schomburgk
has himself described individual species of his collection
(ibid., pp. 12, 375). Gardner has published the diag-
noses of 100 new plants, discovered by himself in Brazil,
as a continuation of his former work (ibid., pp. 97-136).
K. Miiller has resumed the description of Gardner's
Mosses (Bot. Zeit., 1845, pp. 89 et seq.) The continu-
ation of Naudin's contributions to the flora of Brazil
(see the preceding Report) comprises the Melastomacese
(Ann. Sc. nat., iii, pp. ] 69-92, and iv, pp. 48-57).

468 BOTANICAL GEOGRAPHY.

Jameson has described a botanical excursion made at
Chimborazo (Lond. Journ. of Bot., 1845, pp. 378-85).

The aqueous vapours of the winds from the sea are precipitated upon the
west side of the western Cordillera of Ecuador, to which Chimborazo belongs.
Hence, simultaneously with the rainy season of the coast of Guayaquil, wet
weather prevails there from the end of December to the middle of May,
whilst on the eastern slope, and on the elevated surface of Biobamba, the
weather is fine. This contrast exerts an important influence upon the vege-
tation ; hence the numerous Calceolarias and the Alstra3inerias are confined to
the western slope ; hence also, in the upper regions, tall-stemmed woody
plants are isohypsilous with the shrubs of the central Cordillera. Between
13,000' and 14,000' Polylepis lanuginosa, one of the Sanguisorbese, forms a
distinct woody zone, regarding which Jameson remarks, that this tree will
grow at a higher level than any other upon the globe. Lower down, on the
road from Riobamba to the locality of Guaranda, which is situated on the
western side of the Chimborazo chain, there exists a meadow-region of the
same extent, until, at 12,000', woods of Aristotelia Maqui and Columettia
sericea are again met with, in which the underwood consists of shrubby
Synantheracea3, Rosaceae, Melastomacese, and Scrophulariaceae. The report
concludes with a list of the families of plants which were found to occur be-
tween 12,000' and 14,000'. Nearly 250 species observed there by Jameson
belong to about 50 families. Those containing most species are: 29
Synantheraceae, 15 Scrophulariaceae, 11 Graminaceae, 11 Rosaceae, 8 Legu-
minosae, 7 Gentianaceae, 7 Umbellifera3, and 7 Cruciferae; 14 Ferns and 13
Mosses; also the following characteristic alpine forms : Ranunculacese (5)>
Caryophyllaceae (4), Ericaceae (4), Vacciniee (3), Valerianacese (4), Orchidacese
(5), and Cyperacese (3). South American forms : Loasese (2), Passiflorese
(1), Escallonia (1), Columellia (1), Solanaceae (5), and Lobeliaceae (2). The
following tropical forms are also found at this level : Melastomaceae (4),
Homaliacese (1), Loranthaceae (2), and Bromeliaceae (2).

Bridges has reported upon the first-fruits of his bota-
nical travels in Bolivia (ibid., p. 571). The first part of
the botanical division of a very important work upon
Chili, by Cl. Gay, has reached us (Historia Fisica y
Politica de Chile, por Cl. Gay. Botanica, torn, i,
pp. 1-104. Paris, 1845, 8vo). The diagnoses are in
Latin, the descriptions in Spanish. The prodromus, when
completed, will contain all the plants of Chili, and a select
number will be illustrated by copper-plate engravings ;
but it also includes garden plants.

BOTANICAL GEOGRAPHY. 469

The genera treated of in the first part, the indigenous ones, are the follow-
ing : Ranunculacese : Anemone, 7', Hamadryas, 2 ; Barneoudia, Ranunculus, 18 ;
Psychrophila, 4 ; andPceonia. Magnoliacese : Drymis, 2. Anonacese : Anona, 1-
Lardizabaleoe : Lardizabala, 2 ; and Boguila, 1. Berberidese : Berber is, 23.
Papaveracese : Argemone, 3 ; and Papaver and Fumaria, 1.

V._ AUSTRALIA AND THE SOUTH-SEA
ISLANDS.

J. D. Hooker opposes the opinion that all or most of
the South-Sea Islands belong to the same primitive
formation (Lond. Journ. of Bot., 1845, p. 642).

The resemblance of their vegetation is rather apparent than real, and is
principally evidenced in littoral plants, and in those which with man have
migrated beyond their native country towards the East. However, that the
original vegetation, with which those naturalized have become associated, is
endemic to the larger groups of islands, at least, is shown by a comparison of
the flora of the Sandwich and Society Islands, for instance, both of which
are subject to the same climatal conditions ; one being situated north, and the
other south of the equator. Tew only of the prominent genera are found in
both groups. The Society Islands are the poorest, but tropical in their
forms, and less peculiar : here the extensive families of the torrid zone pre-
dominate, as the Malvacese, Leguminosse, Apocynese, Urticacese, Melasto-
macea3, and the Myrtacese. Of the forms peculiar to the Sandwich Islands,
the Synantheracese, Lobeliacese, Goodenoviacese, and Cyrtaridracese, few or
no representatives are found. Other families, as the Grasses, Euphorbiacese,
Rubiacese, &c., which are numerous in both Archipelagos, occur for the most
part in isolated species. The same view of the endemic character of the
Flora of the Sandwich Islands is adopted by Hinds. (Ann. Nat. Hist, xv,
pp. 91-3.) With other Floras and those of the most different kinds, isolated
points of resemblance only can be shown. Of 165 species which the traveller
collected upon the coast there, half are endemic. In a physiognomical point
of view the amount of forests is small in comparison with that of other tro-
pical countries, the trees are not tall and only crowded in moist sheltered
valleys. Cinchonaceae, Guttiferae, Sapindacese, Euphorbiaceae, are found there,
mixed with Tree-Ferns, and a single Palm which was originally endemic.

The work of Strzelecki upon New Holland contains a
number of valuable details on the conditions of the vege-
tation of this continent (Physical Description of New South
Wales and Van Diemen's Land. London, 1845, 8vo).

The extra-tropical south-eastern coast is exposed pretty uniformly to

470 BOTANICAL GEOGRAPHY.

variable winds, which are dependent upon the monsoons of the adjacent
oceans, but are not the same in the different latitudes. At Port Jackson
and Port Macquarie (32 S. lat.) equatorial winds prevail in the summer and
polar currents in the winter ; in Port Philip (south-eastern extremity of the
continent) equatorial currents in the winter, and polar currents in the sum-
mer ; in Van Diemen's Land the equatorial winds predominate throughout the
whole year (p. 168). The amount of rain is far more considerable on the
coast than we should expect : on the average it amounts to 48" 6 in New
South Wales, and to 41" 3 in Van Diemen's Land (p. 192). The temperature
is far more uniform than in corresponding latitudes of the northern hemi-
sphere, as is shown by the following table (p. 229).

Woolnorth in

Port Port Port Van Diemen's

Macquarie. Jackson. Philip. Laud.

Mean temperature +20 c. -f!92c. +16 3 c. -f!4lc.

+ 23 9 " +23 2" +20 8" +16

Mean winter tern- ) +16 o r +15 i>/ +11 9" +12 3

. perature 5

Maximum of summer +31 3"* 4-27 8"f + 32 5f +204J

winter + 8 2* + 7 4 + 27f + 8J

The influence of the geological conditions upon the vegetation and the cul-
tivation of the soil is exceedingly variable, according to Strzelecki, as shown
by a comparison of New South Wales with Van Diemen's Land. In New
South Wales, granite, sandstone, and conglomerate predominate ; limestone
is confined to but few localities ; in Van Diemen's Land, porphyry, greenstone,
basalt, and trachyte predominate ; limestone is also common (p. 360). In
the former, the silica existing in the soil favours the nocturnal diminution
of temperature, and would act still more prejudicially if the vegetation, which
is more dense, did not frequently give rise to the formation of clouds (p. 219).
But the small quantity of soluble constituents in the soil renders it only
adapted for indigenous plants, as for pasture-land, but not for agriculture.

The botanical letters from New Holland, by Leichardt
(Lond. Jonrn. of Bot., 1845, pp. 278-291), whose great
voyage of discovery through the interior of the continent,
which has never been surpassed in its results, was de-
scribed without a view to publication, excite the most
sanguine hope that the botanical characteristics of
Australia, taken up by such talent for observation, and
described in an equally successful manner, will, at some

* The warmest month is November ; the coldest, August.
f The warmest month is November ; the coldest, July.
| The warmest month is January ; the coldest, August.

BOTANICAL GEOGRAPHY. 471

future time, acquire important elucidations from this
traveller.

Systematic contributions to the Flora of Australia :
Bonder's diagnoses of 76 new Algse, belonging to Preiss's
collection from the Swan River (Bot. Zeit., 1845,
pp. 49-57) Berkeley's new Fungi (54 sp.), from the
same locality, and belonging to Drummond's collection.

J. D. Hooker has written a memoir upon the distri-
bution of the Coniferae in the southern hemisphere (Lond.
Journ. of Bot., 1845, pp. 137-157).

Van Diemen's Land contains ten different Coniferse, which are endemic to
the island. They occur in limited localities, and most of them were dis-
covered by Gunn ; they are Callitris australis Br. (Oyster-bay pine), a tree
from 50' 70' in height ; C. Gunnii J. D. Hook, (native cypress), 6' 10' in
height ; Arthrotaxis, 3 sp. ; Micocachrys tetragona J. D. Hook., a tree from
15' 20' high ; Podocarpus alpina Br., a shrub on Mount Wellington, at an
altitude of from 3' 4000' ; Podocarpus Lawrencii J. D. Hook. ; Phyllocladus
asplenifolia Rich, (celery-topped pine), 50' 60' high ; Dacrydium Franklinii
J. D. Hook. (Huon-pine), the most beautiful tree of them all, from 60' to 3 00'
in height, with a diameter of from 2' 8', of limited occurrence, but used at
the harbour of Macquarie as ship-timber.

Sketch of the distribution of the Coniferse as yet discovered in the southern
hemisphere : 16 species in New Holland (10 Callitris, 4 Podocarpus and 2
Arauearia at Moreton Bay), 10 species in Tasmania (vid. sup.), 13 species
in New Zealand and the South Sea Islands (6 species of Podocarpus, of
which the Kaikatia, Podocarpus dacrydioides Rich, is most common at the
Bay of Islands, 3 Dacrydium, Thuja Doniana Hook., Phyllocladus tricho-
manoides Don., Dammara Australis = Kauri pine, Arauearia excelsa Ait.
=Norfolk Island pine, probably confined to this island ; 8 species in South
America (4 Podocarpus in Chili and the Brazils, Thuja chilensis Hook. ;
andina Popp. ; Thuja tetragona Hook.= Alerse of Chiloe, Arauearia bra-
sz'/^Tm's Brazilian pine, Arauearia imbricata = Chili pine, on the Andes,
from 37 to 46 S. lat. ; Juniperus uvifera Don., from Cape Horn, remains
doubtful ; about 6 species in the South of Africa and the Mauritius (2 Podo-
carpus, 3 Pachylepis, including Pachylepis Commersoni from the Mauritius,
and Juniperus capensis Lam., doubtful.

We have now received 15 Parts of J. D. Hooker's
illustrated work upon his Antarctic Voyage. (The Botany
of the Antarctic Voyage. London, 1845, 4to.)

The character of the vegetation of Lord Auckland's islands is more clearly
described than before (Ann. Hep. for 1843). It was previously mentioned
that these islands, the volcanic soil of which ascends in the form of gentle

472 BOTANICAL GEOGRAPHY.

hills to an elevation of 1500', were uniformly covered with forest-, shrub-,
and pasture-land. The forest on the abundant humous soil of the coast con-
sists of Metrosideros lucida, mixed with an arborescent Dracophyllum, the
underwood consisting of Coprosma, one of the Rubiacese, shrubs of Veronica
and Panax. As in New Zealand, beneath the woody plants, social Ferns
are abundant. One of them, Aspidium venustum, Hombr. Jacquin., the
luxuriant foliage of which spreads out from the summit of a stem from 2' to
4' in height, and 6" in diameter, reminding us, in its growth, of the climate
of the Tree-ferns of New Zealand, just as the Dwarf-palm does of a tropical
climate. Above the forest region, which is confined to the coast, the bush-
land alone is found as far as an elevation of 800', where it is gradually
replaced by treeless pastures of herbaceous plants and grasses. The herba-
ceous plants display flowers equalling alpine plants in brilliancy of colour,
and for the most part are vicarious species of alpine types of plants, as
Gentiana, Veronica, Cardamine, and Ranunculus.

Campbell's Island is girdled with rocks, like St. Helena, and does not
therefore contain a connected forest-region. Being covered internally by
meadows, it only contains the Ferns found beneath the bushes in the Auck-
land Islands, in isolated sheltered localities. Of the antarctic forms, a large
golden-yellow Liliaceous plant (Chrysobactron) is found on the rocky heights,
in such luxuriance, that the colour of its flowers is perceived by those
sailing by at the distance of a mile from the coast.

Summary of the Flora of the Lord Auckland's Islands and Campbell's
Island : 3 Ranunculacese (Ranunculus), 4 Cruciferse (Cardamine), 4 Caryo-
phyllaceae (Stellaria, 3 Colobanthus), 1 Drosera, 1 Geranium, 3 Rosacese
(Sieversia and 2 Accend), 3 Epilobium, 1 Callitriche, 1 Metrosideros, 1 Mon-
tia, 1 Bulliardia ; 3 Umbelliferse (1 Pozoa and 2 Anistome*), 1 Panax, 1
Aralia ; 7 Rubiacese (6 Coprosma and Nertera) ; 11 Synantheraceae (Trin-
curon, Ceratella, 3 Leptinetta, Ozothamnus, Helichrysum, 2 Pleurophyttum,
Celmisia, and Gnaphalium), 3 Stylidiacese (2 Dracophyllum and Forstera}, 1 of
the Lobeliacea? (Pratia), 1 of the Epacridacea (Androstoma), 1 of the Myr-
sinacese (Suttonia), 2 Gentiana, 2 Myosotis, 3 Veronica, 2 Plantago, Rumex, 1 ;
Urtica, 2 ; 8 Orchidacese (2 Thelimitra, 2 Caladenia, Chiloglottis, Acianthus,
and 2 undetermined), 2 Asphodeleae (Chrysobactron and Astelia), 5 Junceae
(2 Juncus, 2 Rostkovia, and Luzula), 1 of the Restiacese (Gaimardia}, 6
Cyperacea? (3 Carex, Uncinia, Isolepis, and Oreobolus), 14 Graminacese (2
Hierochloe, 4 Agrostis, Trisetum, Bromus, 2 Festuca, 3 Poa, and Catabrosa),
17 Ferns (5 Hymenophyllum, Aspidium, 3 Asplenium, Pteris, 2 Lomaria, 2
Polypodium, Phymatodes, Grammitis, and Schizeea) ; 66 Mosses, described in
connexion with Wilson ; 85 Hepaticee, described by J. D. Hooker and Taylor ;
30 Lichens, by the same ; 57 Algse, by J. D. Hooker and Harvey ; and 15
Fungi, by Berkeley. Several of the Cryptogamic species are European,
but few only of the Phanerogamia, which have either been introduced, or,
forming varieties, their determination appeared doubtful.

BOTANICAL GEOGRAPHY. 473

The Flora of the Antarctic countries commences with the eleventh part of
the work, and includes all latitudes situated between 45 and 64 S. lat. ;
comprising the following points, which were explored by the traveller,
Fuegia, the south-west of Patagonia, the Falkland Islands, Palmer's Land,
and Kerguelen's Land. Summary of the families as yet treated of : Ranun-
culacese (Anemone, 8 Ranunculus, 3 Hamadryas, and 3 Caltha), 1 of the
Magnoliacese (Drimys), 3 Berberis, 11 Cruciferse (Arabis, 2 Cardamine, 3
Draba, Pringlea antiscorbiitica = i\\Q cabbage of Kerguelen's Land, see Ann.
Rep. for 1843, Thlaspi, Senebiera, 2 Sisymbrium"), 1 of the Bixacere (Azara,
in South Chili), 4 Viola, 1 Drosera, 13 Caryophyllacese (Lychnis, Sagina,
4 Colobanthus, 4 Stellaria, Arenaria, 2 Cerastium), 4 Geranium, 2 Oxalis,
2 Celastrinese in Fuegia (Maytenus and Myginda), 1 of the Rhamnese from
the same place (Colletid), 8 Leguminosee (2 Adesmia, 3 Vicia, and 3
Lathyrus], 15 Rosaceee (2 Geum, Rubus, Fragaria, Potentilla, and 10 Acana),
2 Onagrariae (Fuchsia in Fuegia, and Epilobium), 6 Haloragese (Nyriophyl-
lum, Hippuris, Callitriche, and 3 Gunnera), 5 Myrtacese (Metrosideros in the
Chonos Islands, 2 Myrtus, and 2 Eugenia), 1 Montia, 1 Bulliarda, 1 Ribes, 8
Saxifragete (2 jEscallonia, Cornidia, 2 Saxifraga, 2 Chrysosplenium, and
Donatia). The Umbelliferse are not yet completed.

The description of the Antarctic Cryptogamia in the ' London Journal of
Botany' (see the preceding Ann. Rep.) has been continued ; 38 new Hepaticse
have been published by J. D. Hooker and Taylor (1845, p. 79-97), 76 new
Algse by J. D. Hooker and Harvey (p. 249 to 276, and 293 to 298), who have
also enumerated the Algae of New Zealand, at present 124 species (p. 521
to 551).

The first volume of the botanical text of the illustrated
work, founded upon Dumont d'Urville's antarctic voyage,
has now appeared ; it contains the Cellular Plants, by
Montagne. (Voyage au Pole Sud et dans 1'Oceanie sur
les corvettes Astrolabe and Zelee. Botanique, t. i, Plantes
cellulaires. Paris, 1845, 8vo.)

They consist altogether of 138 Algse, 42 Lichens, 48 Hepaticae, and 40
Mosses. The preface contains lists of the Cryptogamia found in both hemi.
spheres, between the pole and the 50th parallel (they consist of 9 Algae, 66
Lichens, 11 Hepaticae, and 14 Mosses) ; also a list of those species which
occur both in high and tropical latitudes (171 species) ; and lastly, of cos.
mopolite species (8 Algae, 6 Lichens, 5 Hepaticse, and 10 Jungermannise).
Montague's new genera had been previously described in a preliminary work.
The copper-plates belonging to the Cryptogamic section, by Hombron and
Jacquinot, the text of which has not at present appeared, although exquisitely
drawn, have been severely criticised by the younger Hooker (Lond. Journal
of Botany, 1845, p. 28).

474 SYSTEMATIC BOTANY.

^.SYSTEMATIC BOTANY.

IN conformity with the character of the earlier sys-
tematic literature, the description of new forms still pre-
dominates, so that even those most capable of the task
are still too much drawn away from the more profound
establishment of the System of Plants. But as in this
Report the latter direction will principally be kept in view,
its brevity will not only be excusable in consequence of a
defective knowledge of the literature, of which important
papers often reach me too late, but will also be an inten-
tional result of the plan of the work.

In January, 1845, the ninth volume of De Candolle's
ProdromusSystematisNaturalis(Paris,8vo) was published,
the tenth followed it in April 1846. The families treated
of will be mentioned presently. Of Walper's Reper-
torium of the diagnoses contained in recent botanical
works, (Repertorium Botanices Systematise, Lips.,
1845-6, 8vo,) the conclusion of the Labiatse appeared in
the last part of the third volume ; in the fourth volume,
which has not been continued beyond the first fasciculus,
the Verbenacese, Myoporinaceae, Selaginacese, Globula-
riaceae, and Plantaginacese, and in the fifth volume, sup-
plements to the Polypetalous families treated of in the
first volume, principally consisting of a reprint of Jussieu's
Monograph of the Malpighiacese ; these extracts and re-
prints are, however, any thing but accurate, as is well known.

A new part of Sir W. Hooker's ' Icones Plantarum,'
containing 50 plates, has been published. (Part 15,
vol. viii, p. 1, Nos. 701-750. London, 1845, 8vo.)

Leguminosee. Bentham is describing the Mimosese, and has given a com-
plete synopsis of the genera and species of this group (Lond. Journal of
Botany, 1844-5) ; during the past year, only Inga, with 134 species. He
reduces this genus within narrower limits ( = Euinc/a Endl.), and remarks
that either the Monadelphous Mmioseae, i. e. one third of all which are
known, must be arranged in a single genus, or the formation of the leaves

SYSTEMATIC BOTANY. 475

must be recognised as a generic character. Thus he separates Inga from
Picetholobium (which has doubly pinnate leaves) by the simple pinnation only ;
but in this manner also obtains habitual characters in the more elongated
and pubescent flowers, and in the thicker legumes, which are tumid at the
margin. It must undoubtedly be admitted as a correct maxim, that when
the higher systematic sections, as the families, are limited by the characters
of their vegetative parts, the inferior categories, such as the tribes and
genera, may also be made to depend upon them, when a natural sub-
division of the group is effected by that means. Alexandra, the new
genius of the Sophoreae, a tree from British Guiana, with colossal flowers,
has been described by Rob. Schomburgk (1. c. 1845, p. 12). The revision of
the genus Genista, by Spach (Ann. Sc. Nat., iii ser. vol. 2, 3), contains a
considerable number of new species ; but, like the previous systematic works
by this author, it cannot be regarded as conclusive, or as a description of
the matter contained in it corresponding with the genius of science, but
merely an excessively prolix enumeration of descriptive details. Part of the
new species consists of unimportant forms, as evidenced by the description
of several belonging to Genista tinctoria ; the diagnoses, of excessive, totally
unnecessary length, do not by any means afford a synopsis of the distinctive
characters, but rather defeat their object, and, mingled with the extended
yet special and abbreviated descriptions, which do not facilitate the recog-
nition of the species as such, since they combine variable with constant
characters, necessarily render this more difficult. The arrangement of the
sections and subgenera is of more importance ; they are also unnecessarily
increased ; but they contain analytical details and new observations, which
will be of use for a future monograph. Dendrosparton Sp. (iii, p, 152) =
Spartium aetnense Biv., and Gonocytisus Sp. (p. 153)= Sp. angulatnm L., are
separated from Genista, and placed in distinct genera.

MYRTACE^E. J. D. Hooker and Harvey have described Backhousia, a new
genus from New South Wales (Bot. Mag. 1845, i, 4133).

MELASTOMACE^E. Naudin separates Microlicia alsinefolia D. C. and
variabilis Mart, from Microlicia, on account of the somewhat different struc-
ture of their anthers, forming them into the new genus Uranthera, and retains
Chatostoma D. C., the proposed character, however, does not contain any
characters distinctive from Microlicia (Ann. Sc. Nat. iii, 3, p. 189, 190).
He elevates Arthrostemma sect. Monochatum into a separate genus, under the
name of the section (4, p. 48). New genera : Octomeris Naud., a shrub be-
longing to the Andes, to which Mel. octona Humb. Bonpl. also belongs
(p. 52) ; Stephanotrichum Naud. (p. 54), and CMloporus Naud. (p. 57), both
from New Granada.

LYTHRARTE^;. Planchon (Lond. Journ. of Botany, 1845, p. 474) refers
Henslovia Wall, (one of the Hensloviaceae Lindl.) to this order. He
ascribes to this genus a Capsula loculicida, valvis media septiferis basi et apice

476 SYSTEMATIC BOTANY.

connexis> and places it near Abatia K P. From the figure given in the
' Flora Peruviana,' he also regards Alzatea R. P. (Celastrinea dulia) as be-
longing to the Lythrarieae, and places Crypteronia Bl. (Rhamnea dub. Endl.)
and Quilanum Blanc, (dub. sedis Endl.) as doubtful synouymes of Henslovia ;
he is, however, only guided by the descriptions of the plants.

DIOSME^E. To this order Planclion refers a dioecious genus of woody
plants from the Malay Archipelago, which he describes as Rabelaisia n. gen.,
without, however, being acquainted with the structure of the ovary (1. c.
p. 519). At the same time the author proposes some changes in the limita-
tion of the Diosmeae, with which be considers the Zanthoxylaceae ought to
be united, after having separated Brucea and Ailanthus from the latter group,
as proposed by Bennet, and in conjunction with Soulamea (Cardiophom
Benth. according to the author's dissections), which has hitherto been placed
among the anomalous Polygaleae, combined them with the Simarubaceae.
Torrey and Fremont have described a new genus Thamnosma, from Upper
California, nearly related to Zanthoxylon (Frem. Exploring Expedit. Americ.
edit, from the Bot. Zeit. 1847, p. 141).

OCHNACEJE. Sir W. Hooker refers Hostmannia n. gen. (Hook. ic. t. 709)
from Surinam, to this family, notwithstanding its bilocular ovary.

EUPHOKBIACEJE. Planclion has described 2 Australian genera (1. c. p. 471,
t. 35, 16): Stachystemon'Pl., which is nearly related to Pseudanthus, and
Bertya PI. to Calyptostigma.

SAPINDACEJE. Snake-seed, which has lately been introduced into com-
merce, consists of the spiral embryos, divested of the testa, of Ophiocaryon
Schomb., one of the Sapindaceae, the snake-nut tree of Essequibo, formerly
referred by its discoverer, Rob. Schomburgk, to the Anacardiaceae, but which
he has now described more completely, and placed in its proper family (1. c.
p. 375-8).

MALVACEJS. Duchartre has published an important account of his re-
searches upon the development of the flowers of the Malvaceae (Ann. Sc. Nat.
iii, p. 123-50), upon the merit of which Ad. Jussieu has expressed himself at
length (Compt. rendus, 1845, Aug. p. 417-26). The outer calyx, at the
period of its earliest formation, appears to represent a bracteal system.
Duchartre considers that the synsepalous calyx is formed in the same manner
as all monophyllous floral envelopes, not by the growing together of originally
distinct organs, as Schleiden believes, but the tube of the calyx is first
formed, from the upper margin of which five sepals spring up. According to
my more recent investigations, which were principally made upon the calyx
of the Onagrariae, this view is essentially in accordance with nature ; but the
sequence of the phenomena is incorrectly described. The free apices of the
organ are first formed, the basilar formative points then unite, in conse-
quence of the lateral growth of each individually, and thus, after the formation
of the lobes, a connected tube of the calyx springs up from the torus. The

SYSTEMATIC BOTANY. 477

marginal union of floral organs of the same whorl, when occurring, must be
considered merely as an exception, in opposition to the universality of this
process. Puchartre's most important discovery relates to the position of the
stamens, and serves to confirm the supposed affinity of the Malvaceae to
the Rhamnaceae. After the formation of the calyx, the stamens are de-
veloped somewhat before the corolla (as in several families with the stamens
opposite), as five rudiments of leaves (mamelons) which alternate with the
segments of the calyx. These divide while their formation is yet scarcely
completed, at first into two segments (dedoublement collateral), in the same
manner as a divided leaf (their development proceeding more rapidly at the
two sides than in the median line, the five primitive eminences become five
pairs of minute rounded tubercles). Almost simultaneously with the division
of those stamens which are first formed, the petals appear; they are opposite
the former, and a considerable distance apart. The polyandrous character
is produced by the frequent repetition of the same formation in front of the
above ten stamens, united in pairs, that is, on their inner side (parallel de-
duplication : five new pairs of tubercles are formed in a circle, which is
situated more internal and opposite to the first). This multiplication of the
stamens is not considered by Duchartre as arising from the formation of
new and opposite whorls upon the torus ; but he appears to regard them,
and certainly with truth, as formed from the expansion of the substance of
the primary leaf toward the interior. The Polyandrous character is often
increased by a second collateral division of the individual stamens. In fact,
in Malope trifida y and some other species, Duchartre has even finally observed
a third collateral division, both of the anthers and of the stamens, so that
here, and perhaps generally, the unilocular anthers ought to be considered as
the halves of a truly dimidiated stamen. Five teeth to the tube of the fila-
ments, which alternate with the petals, appear to be universal in the buds,
and are considered as forming a second circle of stamens, without any con-
vincing argument being brought forward. Regarding the pistil of the Mal-
vaceae, Duchartre assumes the existence of five primary forms, the two
former of which agree in the circumstance, that at first a pentagonal collar-like
protuberance (bourrelet pentagonal) arises from the torus at the circum-
ference of the apex of the axis (mammelon central), the angles of which are
opposite the getals (at least this position is -mentioned as occurring in
Malope} ; either numerous carpels then shoot out from the margin of this
protuberance (Malopeae), or five only from its angles (Hibisceae). The
formation of the carpels is also preceded in the Malveae and Sideae
by a protuberance, which is not, however, pentagonal, but annular. The
number of carpels growing from its margin is undetermined. Lastly, the
greatest deviations occur in Pavonia, and some allied genera, in which, upon
an annular protuberance, the rudiments of ten styles arc said to appear first,
and subsequently to be fused into five ovaries.

478 SYSTEMATIC BOTANY.

HYPEEICACE^:. Cossoii and Germain (Flore de Paris) admit Spach's
genus Elodea (Hypericum elodes), which differs from Hypericum in its parietal
placentation, whilst the latter has a central placenta. To me, however, the
difference appears merely to consist in the parietal placentae of Hypericum
meeting in the axis of the fruit, whilst in Elodea this is not the case : whether
this is a generic character or not must be decided by a future monograph of
the family, Spach's work not being sufficient for this purpose.

CARYOPHYLLACE.E. J. Gay's monograph of Holosteum (Ann. Sc. Nat. iii,
4, p. 23-44) is characterised by the author's well-known accuracy ; but is
impaired by the prolixity which is unfortunately so frequently combined with
such accuracy, particularly with endless quotations. The following new
genera are proposed by Gay in this memoir : Rhodalsine G. (p. 25) =
Arenaria procumbens V. It appears to differ from all the other Alsinese, in
the stamens being biserial, which is, however, a very relative character only :
and Greniera (p. 27) = Alsine Douglasii Fzl. and Arenaria tenella Nutt. ;
characterised by its seeds, which are compressed discoidally.

CACTACE^E. We are indebted, for a scientific summary of the Cactaceae,
to the Prince Salm-Dyck, who possesses the largest collection in existence
on the Continent (about 700 forms), and is also one of those best acquainted
with this difficult botanical group (Cactea3 in horto Dyckensi cultse, additis
tribuum generumque characteribus emendatis a principe Jos. de Salm-Dyck
Paris, 1845, 8vo). Pfeiffera S. (p. 40), is a new genus described in this work.

CUCURBITACEJE. Wight has written in the ' Madras Journal of Science,'
and Gardner in the 'Lond. Journ. of Bot.' (1845, p. 401), in favour of
Seriuge De Candolle's view, that the middle of the carpels is situated in the
axis of the fruit, and that the cells of the fruit are formed by the revolute
incurvation of their margins, and have endeavoured to support this para-
doxical theory by the course of development of the ovary. According to
Gardner, the external wall of the fruit is formed by the tube of the calyx
only, with which, in Coccinia indica, the dissepiments are merely loosely
in contact, without adhering to it. The course of the bundles of vascular
tissue also, the principal trunks of which, both in this plant and in Bryonia,
are situated in the axis, is in favour of Seringe's view; but the principal
point in the solution of this question consists in distinguishing the placentae
from true carpels, which has not yet been accomplished. It is still extremely
improbable that three leaves should grow out of the point of the axis.
Payer remarks (Ann. Sc. Nat., iii, 3, p. 163) that at the lower joints, where
three vascular bundles enter the petiole, the stem of the Cucurbitaceae is
not furnished with tendrils, whilst on the other hand, in the case of the
upper leaves, according as one or two tendrils are present, it receives two
vascular bundles only, or the central one alone. He thus explains the
oblique position of the axillary bud, which is always situated opposite the
central vascular bundle, and thus, where as usual a single tendril only ac-

SYSTEMATIC BOTANY. 479

companies the leaf, obtains an oblique position. But he does not thus prove
that the tendrils are segments of a leaf or stipules, whilst if they are re-
garded as entire leaves, this may be proved by the earlier stage of develop-
ment, before the formation of any vessels (Wiegm. Archiv., 1846, p. 24).

CRUCIFEILE. Barneoud has described the small group of the Schizope-
talese, to which, in addition to the principal genus (containing 2 species),
Perreymondia, n. gen. Barn, from Chili (with 4 species) belongs (Ann. Sc.
Nat. iii, 3, p. 165-8). The characters are limited to the divided petals and
the branched hairs, Perreymotidia not possessing the divided cotyledons,
but an ordinary notorrhizal embryo, and as this is the only difference, it can-
not constitute a distinct genus. Trautvetter separates Matthiola deflexa, Bg.
from the genus Matthiola, as Microstigma, Tr. (PI. Ross, imagines T. 25).
Nejw genera : Lyrocarpa, Hook. Harvey (Lond. Journ. of Bot. 1845, p. 76) ;
it has a panduriform silicule, and was discovered by Coulter in California ;
Dithyrea, Harv. (1. c. p. 77), allied to Biscutella, from the same source ;
Oxystylis, Torr. Erem. (Explor. Exp. and 1. c. p. 41), well characterised,
approaching the Capparidacesc, also from California ; Pringlea, Anders, d.
Hook. (Antarct. Voy. p. 238, pi. 90-1), the above-mentioned cabbage of
Kerguelen's Land.

PAPAVERACEJE. New genera from California : Romneya, Harv. (1. c. p. 73),
principally distinguished from Papaver by the trimerism of the two outer
whorls ; Arctomecon, Torr. Erem. (1. c. p. 40), only differing from Papaver,
according to the description, in the strophiolate seeds.

RANUNCULACE.E. Eor a notice of Barneoud' s work, the whole of which
has not yet been published (Compt. rend. 1845, ii. p. 352-4), see Link's
' Physiological Report ' (p. 95). Cl. Gay has established two genera from
Chili : Psychrophila (Hist, de Chile Bot. i. p. 47, t. 2), separated from
Caltha and Barneoudia (ib. p. 29, t. 1, f. 2), allied to Hetteborm.

SAXIFRAGACE^E. Gardner describes a shrub which he discovered on the
Organ mountains at Rio, as Raleighia (Lond. Journ. of Bot. 1845, p. 97),
with the following essential characters : Calyx four-partite, valvular, corolla
absent, stamens numerous and perigynous, ovary one-celled, with a single
style and 3 ( 2) placentae, supporting numerous ovules, which are sub-
sequently situated on the median line of the valves of the capsule ; seeds
with an axile embryo ; leaves opposite, connate at the base, and serrated.
The author refers it to the Bixacese, but Bentham rightly places it in the
Cunoniaceee near Belangera, as it forms a transition link from this to the
parietal families, by its truly parietal placentae, but differs from the latter
by their attachment. Planchon takes a different view of Raleighia
(ib. p. 476) ; from the examination of dried specimens, he regards this genus
as scarcely generally distinct from Abatia (Lythrarese, vid. Sup.), which
cannot be the case unless both Gardner and Bentham have described the
fruit and seeds totally incorrectly.

480 SYSTEMATIC BOTANY.

UMBELLIFEBJE. New genus from Lord Auckland's Archipelago : Aniso-
tome J. D. Hook. (Antarct. Voy. p. 76, t. 8-10). The Umbellifene, Crassu-
lacese, &c. proposed in the ' Phytographia Canariensis,' will be passed over
until the work is completed.

EPACBIDACE^E. New genus : Androstoma J. D. Hook., from the Auckland
Islands (Autarct. Yoy. p. 44, t. 30).

MYRSINACE^E. New genus: Labisia, Lindl. (Bot. Reg. 1845, t. 48),
from Penang, differing in the induplicate aestivation of the corolla.

BiGNcraiACEJE. In the Prodromus, this family is treated of by De
Candolle, jun., in conjunction with the Sesamese (vol. ix), in accordance with
the description prepared by De Candolle, sen. The Sesamese, which in this
work also include the Pedalinese, appear only to be separated from the
Bignoniacese, because the type of the fruit is assumed to be quinary. The
African species are separated from Sesamum, as Sesamopteris. The fol-
lowing genera are distinguished from Bignonia : Pachyptera,
=B, uncinata Mey., Anemoptegma Mart., Distictis Mart.,
Mart., Cybistax Mart., Adenocalymna Mart., Sparratosperma Mart., Hete-
rophragma = B. quadrilocularis Roxb., Craterocoma Mart. The Cres-
centieae, which are arranged by Endlicher among the Gesneriaceae, form, in
this work, the second tribe of the Bignoniaceae, being characterised by the
indehiscent fruit and the wingless seeds, found principally in Madagascar :
Tanaecium pinnatum W. is separated from Tancecium, as Kigelia. Parmentiem
is a new genus from Mexico. The position of Bravaisa= Bignonia bibrac-
teata Bert, remains doubtful.

GESNERIACEAE. The Gesneriacese having appeared earlier in the Prodro-
mus, the Cyrtandracese form an independent family in the ninth volume, the
elder De Candolle having left them prepared. Ramondia2tfidiHaberlea,w\t\\
Conandron from Japan, are correctly brought into this family as a distinct
group, on account of the septicidal dehiscence of the capsule.

ACANTHACE^E. New genera : Lankesteria Lindl., from Sierra Leone (Bot.
reg. Miscell., 1845, p. 86?; Whitjieldia Hooker, from the same source (Bot.
Mag., p. 4155) ; Salpixanthia Hook., from Jamaica (ibid., p. 4158),

SCROPHULARIACEJS. Bentham's Monograph fills the greater part of the
tenth volume of the Prodromus. With the exception of the Salpiglossidea?,
which, notwithstanding the stamens being anisomerous, would have been
more properly excluded, and placed among the Solanaceae, all the genera
possess an imbricate aestivation of the corolla. The position of the fourth
and fifth petals, which form the upper lip in those furnished with labiate
flowers, separates the tw