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THE ANATOMY OF SOME DESERT PLANTS

BY

AMELIA L. KELLOGG

THESIS

FOR THE

DEGREE OF BACHELOR OF ARTS

IN

SCIENCE

COLLEGE OF LIBERAL ARTS AND SCIENCES

UNIVERSITY OF ILLINOIS

1915

Digitized by the Internet Archive
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UNIVERSITY OF ILLINOIS

THIS IS TO CERTIFY THAT THE THESIS PREPARED UNDER MY SUPERVISION BY

Peer e hier ig mere UO TEM Ncw iy a a

ENTITLED The Anatomy of. Some Desert..Plants. _

IS APPROVED BY ME AS FULFILLING THIS PART OF THE REQUIREMENTS FOR THE

DEGREE OF

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Contents.

A. Introduction.
B. Methods.
C. Anatomical Studies.
I. Chenopodiaceae
1. Atriplex canesens.
(a) Stem.

(b) Leaf.

2. Atriplex Nutallii corrugata.

II. Loaseae.

1.Mentzelia: multiflora.

(a) Stem.

(b) Leaf.

III. Rosaceae.

1.Cercocarpus ledifolius.

Stem
Leaf

D. Plates

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The Anatomy of Some Desert Plants.
The anatomy of relatively fan American plants has been studied in
| detail. Solereder in his very excellent work, " The Systematic Anatomy of
Dicotyledons " , has brought together the chief anatomic features that
characterize the families of dicotyledonous plants and a careful examination
of this work reveals the fact that American botanists have contributed very
little to the wibieets
The greatest variation in anatomic structure is shown in the families
whose representatives possess the widest adaptibility to environmental condi-
tions. Schimper, Warming, Volkens and others in their ecologic studies have
called special attention to changes in anatomic structure induced by environ-
mental conditions, chiefly physical. It is with the view of adding a little
to our knowledge of the anatomy of American plants, particularly such as have
adapted themselvws to extreme conditions, that the present study was under=
taken. The plants studied are :
(1) Atriplex canesens, Wats.
(2) Atriplex Nutalii corrugata, A. Nels.
(3) Mentzelia multiflora, Gray.
(4) Gercocarpus ledifolius, H.B.x.,
of the families Chenopodiaceae, Loaseae, and Rosaceae respectively.
The material was collected in the Colorado Desert, near Palisade,
Colorado, at an elevation of between five and seven thousand feet above sea

level, and preserved in a mixture of the following composition:

850 cc. water,

| 36 cc. formalin
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250 cc. alcohol (ninety-five per cent)

In the preparations of sections suitable for anatomic study, great

difficulties were expsrienced due to the extreme hardness of the more resist-

ant tissues of both leaf and stem. The material embedded by the ordinary

method in paraffin or celloidin was utterly worthless for study. Soaking in

alcohol and glycerine with or without subsequent embedding, gave no better

results.

a
Finally, the method so successfully employed wy Jeffries and Plowman

The preserved material was washed

was tried and found to give good results.

for some time in eighty per cent alcohol, and was then transferred into para-

ffined bottles containing a ten per cent solution of hydrofluric acid. In this

acid, with two changes of solution, it remained for two weeks. The material

was then washed in flowing water to remove the acid, dehydrated by passing

it through solutions of thirty, fifty, seventy, sighty, ninety, ninety-five

per cent, and absolute alcohol respectively, and embedded by the ordinary

process in paraffin or celloidin. The harder, more resistant stems and more

brittle leaves were infiltrated and embedded in celloidin. The infiltration

was accomplished by transferring the tissues successively into two, four, six,

sight, ten, and fourteen per cent sgolutione of celloidin in equal parts of

absolute alcohol and ether. The material remained in the two, four, e#ix,

eight, and ten per cent solutions, respectively,for two days, and in the

fourteen per cent for two wesks. It was then embedded in the usual manner.

The sections were stained in Bismark brown, fuchsin and eosin. In

soms cases, it was found possible to study and draw structure in detail with-

eut previous staining of the sections.

1. Atriplex canesenc, Wats. (Chenopodeaceae)

| 2. Atriplex Nutalii corrugata, A. Nels. (Chenopodeaceae)

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3. Mentzelia multiflora, Gray (Loaseae)

4.@ercocarpus ledifolius, H. B. K. (Rosaceae)

The widely different stem structure in the Chenopodeaceaze led
De Bary’ to classify such stems into three classes as follows:

1. The growth in thickness of the stem is due to the formation of
pericyclic rings or strips of cambium which originate and lose their activity
successively, forming secondary bundles and conjunctive tissue. The success-

ive rings thus formed vary somewhat in regularity and completeness.

2. The growth in thickness following the formation of the ring of

bundles, results from the appearence around the margins of the phloem, of a
ring of interfacicular cambium which forms alternately vascular bundles and
intermediate tissue.

3. In the final type, a combination of the processes 1 and 2 takes
place, that is, the normal cambium and normal secondary thickening appear as
in type 1; this however, soon stops and further growth in thickness is contin-
ued by a new extra fasecular zone of cambium which appears outside the primary
masses of phloem - according to type 2.

Solereder's classification disregards type 3, grouping all stem

structures under the two extreme types described by De Bary?

Atriplex canesens.

The stem structure of Atriplex canesens may be considered as belong-
ing to type 2. The vascular bundles are embedded in prosen chymatous
conjunctive tissue and arranged without definite regularity. The conjunctive
tissue is conspicuous in that it is composed of very thick-walled cells con-
taining simple pits. The arrangement is irregular, and nowhere are clearly
defined medulary rays visible.

Within the vascular cylinder is a rather large pith composed of

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large cells full of protoplasm and so closely crowded as to admit of no
intercellular ieadan.

The primary cortex is composed of parenchyma, irregularly broken
areas of collenchyma, and an abundance of sclerenchymous fibres scattered
along the outer margin. The cortex is bounded by a single layer of thick-
walled epidermal cells.

The surface of the stem is covered with a membranous covering of
collapsed water-storage hairs. These are unicellular, irregular in size
and shape, and epidermal in origin. They are very numerous and, when they
collapse upon maturity, form a dead membranous covering that serves to check
transpiration. The leaf of Atriplex canesens is covered by similar hairs,

G Fie. \-2Plate'l: )

The most striking anatomic character brought out in the cross-
section of the leaf is the absence of the usual spongy parenchyma, the greater
part of the leaf being made up of elengated chlorophyll-bearing cells not
unlike the pallisade cells of ordinary leaves. ( Fig. + Platex ) Very
few intercellular spaces are apparent in the transsection.

The vascular bundles are located centrally within the pallisade
tissue. They are characteristically surrounded by a sheath of large, thick-
walled cells, nearly circular in cross-section. (Fig. Z Platet )

Crystals of oxalate of lime occasionally occur in the form of
cluster crystals in the various tissues excepting only the vascular bundles.

The upper and lower epidermis are exactly alike. Each is composed
of a single row of thick-walled cells without gelatinization or waxy deposit.
In both, stomates are fairly abundant and are unaccompanied by subsidiary
cells.

In:the leaf, as in the stem, protection against excessive transpir-

ation is well provided for by a membranous covering of collapsed hairs,

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previously described in connection with the stem. ( Fig.|-2 Plate T )

Atriplex Nutalii corrugata.
The anatomy of Atriplex Nutalii corrugata, the second species of
Atriplex examined, is so nearly identical with the above described species
as to warrant no further comment on structural details. Both correspond very

Closely te Atriplex Hamilus described by Volkens in his study of Egyptian-

Arabian Desert plants.”

The most important anatomical character of the family of Loaseae
is its hairy covering. The hairs may be described as :

1. simple, unicellular hairs of varisd length whose walls may or
may not be silicified;

2. unicellular, conical, charp-pointed heirs and strongly silici-
fied. The surface is covered with sharp or blunt spines directed eithsr
forward or backward;

3. unicellular, strongly silicified, barbed hairs of variable
length and often comparable to a many-fluked anchor. The stalk is sometimes
covered with recurvsd spines;

4, long, unicellular, hooked, or climbing hairs with a high multi-
cellular base. These bear several sharp hooks at the apex and show active
protoplasmic movement;

5. long, unicellular, slightly silicified, sickle-likxe hairs, with-
out protoplasmic movement;

6. unicellular hairs, slenderly filiform, with local tuber-like
swellings ut the base and sharply attenuated at the apex or apex with barbs;

7. long, strongly silicified, unicellular stinging-hairs with a
multicellular base and frequently containing a yellow fluid which has a very

irritating effect upon animals;

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8. short, simple, multicellular hairs;

9. multicellilar glandular hairs with an uniserate stalk of varied
length and with a unicellular head.

Other characters essential to the Loaseae are hairs that are calci-
fied rather than silicified; also crystolith-like structures for the most part
impregnated with carbonate-of-lime. These usually occur in the unicellular

hairs, excepting only the stinging hairs, and their subsidiary cells.

Mentzelia multiflora,

Mentzélia multiflora possesses the several kinds of hairs described
under types 1, 2, 3, and 6, excepting that they are calcified rather than
Silicified. This is proved by the fact that they withstood most rigid treat+
ment with the hydrofluric acid that was used in the preparation of sections
for study. thers are crystolith-like structures impregnated with carbonate-
ef-lime. These are restricted to the larger unicellular hairs and their

subsidiary cells.

The stem consists of a woody cylinder, including narrow medullary

reys, fairly wide vessels with simple perforations and with bordered pits in
contact with parenchyma as well as the wood prosenchym. The weod prosen-
chyma has fairly wide lumina. (Fig.{ Plate).
ther tissues are a central pith composed of large, somewhat angular

thick-walled cells; a well marked cambium; and a pericycle of large irregularly
shaped parenchyma cells. The epidermal cells vary in size and shape from
the erdinary brick-shaped cell to the more complex hair-like structures,
previously described. (Fig. | Plate’).

The leaf of Mentzelia multiflera is oval in cross-section. Regard-

ing its structure, it may be pointed out that the stomata have no subsidiary

7.
cells, but are surrounded by a variable number of ordinary epidermal cells.
They occur at irregular intervals throughout the ring of epidermis.

There is a rather conspicuous central vein with clearly different~
iated zone of xylem and phloem, - also a network of tracheids running free
in the mesophyl.

The mesophyl consists of a spongy parenchyma surrounding the central
vein, gradually becoming differentiated into pallisade-like cells as it
approaches the epidermis. (Fig. ! Plate iL).

The epidermis of the leaf is similar to that of the stem except

that the cells are larger in proportion and thicker-walled and the hairs are

longer and stronger. Upon the outer surface occurs a triple layer of cutin
not present on the epidermis of the stem. (Fig. | Plate).

Specific anatomical characters are almost entirely lacking in the
Rosaceas,. According to Solereder and others the following are the most
prevalent characters:

1. .the lack of simple uniseriate clothing hairs;

@. the presence of bordered pits in the prosenchymatous ground-

work of wood in almost all woody species.

Cersocarpus ledifolius.

The stem of Caercocarpus ledifolius is typically woody and possesses
bordered pits in the prosenchymatous groundwork of wood, The vascular
cylinder is a fairly wide region consisting of small lumined wood cells and
narrow medullary rays extending as a conspicuous band out through the
rather well preserved phloem, There is a central pith of large circular
heavy-walled cells. The cortex is rather wide and consists of irregularly
shaped parenchyma cells, increasing in size as they near the evidermis. The

epidermis consists of small, circular cells, regular in size and shape, and

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with a thick deposit of cutin on its outer surface. (Fig. 1 Plate W ).

A large amount of tannin and silica, deposited in the cells of

parenchymatous tissues of this stem, gives it a very conspicuous character.

The leaf of this same plant is peculiar in cross-section in that

there is a prominent mid-rib from which the blade curls back on either side,

giving the entire structure an appearance not unlike the figure three. This

mid-rib contains a single vein with a definite xylem and phloem broken by

narrow bands of parenchyma cells. (Fig. | Plate Ww ). Below the mid-vein

is a crescent of spongy parenchyma meeting the small crowded pallisade-like

eslls above - which compriss the bulk of the internal tissues. The lateral

veins proceeding from the mid-rib pass slightly upward and are distributed

Each of the lateral veins is surreunded by a sheath

through the mesophyl.

of heavy-walled colls, one layer deep. (Fig. | Plated ).

A conspicuous layer of hypodermis occurs just below ths epidermis.

It is composed of large thick-walled cells abundantly supplied with tannin.

This is also true of the cells composing the bundle-sheaths.

The epidermis consists of a single layer of very small ce

Stowates are

a well developed deposit of ecutin on the cuter surface.

abundant on the lower surface of the leaf. Transpiration is reduced by a

dense mat of unicellular clothing hairs and the peculiar curling of the leaf.

The hairs are entirely restricted to the under surface cf the leaf on either

side of the prominent mid-rib. (Fig. / Plate Y ).

In the pallisade-like cells of the mesophyll, a considerable

quantity of silica is deposited.

In the study of the above American desert plants, belonging to the

families Chenopodeaceas, Loaseae and Rosaceae, no new or distinct characters,

not already nected, were discovered.

——7~~ ea

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+

References.

‘Jeffries - Plowman: Botanical Gazette, Volume XXXVII

; June 1904, pp.456.
"De Bary: Comparative Anatomy of Phanerogams and Ferns,
pp. 490.
*Solereder: Systematic Studies of Dicotyledons, pp 655.
*Volkens: The Flora of the Egyptian-Arabian Desert.
"Observed by Solereder: The Systematic Study of Dicoty-

ledons, pp.380.

Plate 1. ‘Figure res ea

Wiadie and end sections of the cross-section of

the leaf of Atriplex canesens - showing covering —
of sollansed water-storage hairs - bundle « neat
~ yallieade-like wesophyll, etc. |

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Plate 2. Figure : 1s

Crondeenand of the leaf of lentuelia mitifiom, | ‘the a
showing mid-vein (xylem and phloem) - network of
tracheice running tree in mesophyll = ‘hooked :

trichomes, etc. .

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Plate 3. Figure l.

Cross-section, of the stem of Mentzelia multi-

flora - showing the relative location of tissues.

Plate 3. Figure 2.
Cross-section of the stem ofGercecarpus ledifolius

location of tissues.

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end other most conspicuous characters.

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