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July, August, September, October, November, December, 1910 

Composed and Printed By 

The University of Chicago Frew 

Chicago, Illinois, U.S.A. 


Ratio of phosphate, nitrate, and potassium on 
absorption and growth (with nine figures) 

Oswald Schreiner and /. /. Skinner 

Contributions to the life history of Widdringtonia 

cupressoides (w T ith plates I-III and three 

• figures) ---«.-.- 

Some peculiar fern prothallia. Contributions from 
the Hull Botanical Laboratory 137 (with 
eleven figures) ------ 


Development of the ovulate strobilus and young 
ovule of Zamia floridana (with twenty-two 

Xotes on the sex of the gametophyte of Onocleu 



IF. T. Saxton 31 

Ltd a Pace 49 

Periodicity in Dictyota at Naples (with one figure) /. F. Lewis 59 

The morphology of the Podocarpineae. Con- 
tributions from the Hull Botanical Labora- 
tory 138 (with plates IV-VI) - - - Mary S. Young 81 

The origin of ray tracheids in the Coniferae (with 
sixteen figures) ------ 

On the relationship between the length of the pod • 
and fertility and fecundity in Cercis (with one 

\V. P. Thompson 101 

/. Arthur Harris 117 

W. Bailey 142 


figures)- ------- Frances Grace Smith 128 

Oxidizing enzymes and their relation to "sap 

stain' ' in lumber ------ 

Some effects of a harmful organic soil constituent 

(with eleven figures) - - Oswald^Schreiih 
Some observations on catalase. Contributions 

from the Hull Botanical Laboratory 139 

(with one figure) ------ 

Twin hybrids (laeta and velutina) and their 

anatomical distinctions ----- Frank M. Andrews 193 
An eocene flora in Georgia and the indicated 

physical conditions (with two figures) - - Edward W. Berry 202 

Chas. O. Appleman 182 

David M. Mottier 209 

Relation of soil moisture to desert vegetation 

(with four figures) - Burton Edward Livingston 241 

The effect of longitudinal compression upon the 

production of mechanical tissue in stems 

(with two figures) L. IL Pennington 2 si 




[volume l 


The North American species of Stereocaulon (with 

nine figures) - - - - - - - Lincoln Ware Riddle 285 

The peg of the Cucurbitaceae. Contributions from 

the Hull Botanical Laboratory 140 (with six 

figures) - 

William Crocker, Lee I. Knight, and Edith Roberts 321 

The origin and development of bulbs in the genus 
Eryihronium (with plates VII-X and seven 
figures) -- Frederick H. Blodgett 340 

Reversionary characters of traumatic oak woods 
(with plates XI and XII) . - 

The Pteropsida (with plate XIII) 

Fertilization and embryogeny in Dioon edule. Con- 
tributions from the Hull Botanical Laboratory 

Irving W. Bailey 374 
Edward C. Jeffrey 401 

141 (with plates XIV-XVII) 

Charles J. Chamberlain 4 X 5 

Notes on Chilean fungi. I (with plates XVIII 
and XIX and one figure) - 

The development of the ascocarp of I^eoiia (with 
forty-seven figures) - 

Briefer Articles — 

A modification of a Jung-Thoma sliding micro- 
tome for cutting wood (with one figure) 
An atmograph (with four figures) - 
The sporangium of Lycopodium pithy oides 
(with plate VII) 

The Botanical Congress at Brussels 

Roland Thaxier 43° 

William H. Brown 443 

The epidermal characters of Frenelopsis ramo- 

sissima (with two figures) 
Another new Achlya (with eight figures) 

Some new saprophytic fungi of the Middle 

Rocky Mountain region 

Three interesting species of Claviceps (with 

eight figures) 

Current Literature 

R. B. Thompson 148 
W. L, Eikenberry 214 

Alma G. Stokey 218 

W. G. Farlow and George F. Atkinson 220 

- Edward Wilbur Berry 3°5 

W.C.Coker 3^ 

Ruth Harrison Lovejoy 3%3 

For titles of book reviews see index under 

author's name and reviews 
Papers noticed in "Notes for Students" are 

indexed under author's name and subjects. 

F. L. Stevens and J. G. Hall 460 

- 65, 150, 226, 310, 386, 464 


No. 1, July 14; No. 2, August 18; No. 3, 
No. 5, November 16; No. 6, December 20. 


P. 31, second paragraph, line 4, for 500 read 5000. 

P. 126, line 5 from bottom, for p = 0.500 read r = 0.500. 

P. 214, line 7 from bottom, for b' read b. 

P. 220, line 7, for pod read pad. 

P. 222, line 22, for Linneaus' read Linnaeus'. 

P. 300' line 10 from bottom, for S. glaubescens read S. glaitcescens 

P. 300, last line, for (fig. 8) read (fig. 6). 

P. 318, line 11, for represent read represents. 

P. 355, line 17, for dropper read runner. 

P. 382, line 2, for Alpanes read A planes. 

No. i 




Ratio of Phosphate, Nitrate, and Potassium on Absorp- 

tion and Growth 

Oswald Schreiner and J. J. Skinner 

Contributions to the Life History of Widdringtonia 


W. T. Saxtcn 

Some Peculiar Fern Prothallia 

Lula Pace 

Periodicity in Dictyota at Naples 

I. F. Lewis 

Current Literature 


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XLhc Botanical (3a3ette 

a dfcontblB Journal Embracing all departments of botanical Science 

Edited by John M. Coulter, with the assistance of other members of the botanical staff of the 

University of Chicago. 

Issued July 14, 1910 




GROWTH (with NINE FIGURES). Oswald Schreiner and /. /. Skinner - - - - 


(with plates i-iii and three figures). W. T. Saxton 


SOME PECULIAR FERN PROTHALLIA. Contributions from the Hull Botanical 

Laboratory 137 (with eleven figures). Lula Pace 



PERIODICITY IN DICTYOTA AT NAPLES (with one figure). /. F. Lewis - - 59 








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Botanical Gazette 

JULY igio 



Oswald Schreiner and J. J. Skinner 

(with NINE figures) 

In this paper are reported the results of experiments obtained 
in connection with a study of the effects of harmful soil constituents 
upon plant growth and upon soil solutions and fertilizer action. 
The results here reported are especially with reference to the growth 
made by wheat seedlings in culture solutions containing many differ- 
ent ratios of phosphate, nitrate, and potash, and in regard to the ratio 
of these constituents originally present and removed by the wheat 
seedlings in the course of the experiment. In these investigations 
solution cultures containing the three fertilizer ingredients, namely 
P 2 5 , NH 3 , and K 2 0, as calcium acid phosphate, sodium nitrate, and 
potassium sulfate, respectively, in all possible ratios of one, two, and 
three constituents, varying them in stages of 10 per cent, were prepared, 
the concentration being 80 parts per million in these constituents. 
The selection of the salts as carriers of phosphate, nitrate, and 
potash, and the statement of the results in terms of P 2 O s , NH 3 , and 
K 2 0, are in harmony with the practice in fertilizer work and, for the 
sake of simplicity, these designations are also retained in the present 
paper. The salts selected, it will be seen, are also carriers of calcium, 
of sodium, and of sulfate, and the three salts, therefore, are the best 
that could be selected for giving at the same time other needed 

The culture solutions of these three salts contained in each case 
a total concentration of 80 parts per million of P 2 5 , NH 3 , and K 2 0, 

1 Published by permission of the Secretary of Agriculture. ~~ 


but these varied, as above noted, in the ratio in which they were 
present. Wheat plants were grown in these various cultures and 
observations were made in regard to general development, the effect 
on root growth, and appearance. The green weight of the plants 
was taken at the termination of the experiment. The solutions were 
changed every three days and an analysis made, the phosphate, nitrate, 
and potash being determined, thus giving the concentration of these 
constituents and the ratio existing at the end of every three-day 
period for comparison with the original concentration and ratio. This 
changing of the solutions was kept up for twenty-four days, thus mak- 
ing eight changes. In this work the methods 2 devised in these labora- 
tories for the determination of small amounts of such constituents 
rendered excellent service and point a way for their further use upon 
other problems in connection with the biochemical relationships of 
soils and plants, which have hitherto been impossible of attack. In 
the discussion and presentation of the results, the triangular diagram 
as used in physical chemistry was employed, and has proven very 
useful as a guide in the work for the systematic handling of the experi- 
mental details. The results can best be presented and interpreted 
by its means, and the method should prove very useful as a guide in 
other lines of experimental work where similar relationships are 


The number of solution cultures required in order to have all the 
possible ratios as outlined above is sixty-six. To bear in mind these 
ratios for three different ingredients, together with the ratios of the 
solutions after the plants had grown, and perhaps also the ratios of 
the material removed from the solution, a total of 594 numbers, is 
practically an impossibility, and it is readily seen that in order to 
discuss such a comprehensive experiment as this the material must 
be reduced to a workable basis, so that the various phases of the 
results can be kept in mind and the proper correlation and compari- 
sons made. The triangular diagram as suggested by Schreine- 


2 Schreiner, O., and Failyer, G. H., Colorimetric, turbidity, and titration 
methods used in soil investigations. Bulletin 31, Bureau of Soils, U.S. Dept. Agric. 



macher 3 in 1893 and again by Bancroft 4 in 1902 has been of the 
greatest service to physical chemistry, where both theoretical and 
practical consideration of percentage composition of three component 
parts are concerned. 

In the present investigation we have likewise to consider the three 
component parts of the fertilizer mixture; namely P 2 O s , NH 3 , and 

p 2 o 5 


Fig. i. — Showing the triangular diagram, with the points numbered, which 
represent the 66 culture solutions. 

K 2 0. 


gular diagram 


diagram is shown in fig. 1. It is an equilateral triangle in which the 


3 Schreixemacher, F. A. H., Konzentrierung oder Verdunnung einer Losung 
bei Konstanter Temperatur. Zeit. Phvs. Chem. 11:81. 1893. 

4 Bancroft, \V. D., Synthetic analysis of solid phases. Jour. Phvs. Chem. 
6:178. 1902. 


each of the ingredients, P 2 5 , NH 3 , and K 2 0,as shown in the diagram- 
Each side of the triangle is divided into ten equal parts, and lines are 
drawn connecting these points. 5 In the diagram, for the sake of 
ready reference, the intersections of these lines have been numbered. 
If we consider the line representing the base of the triangle, it is obvious 


that the point which represents ioo per cent K 2 (number 56 in the 
diagram) represents at the same time o per cent NH 3 , and the point 
100 per cent NH 3 (number 66) likewise represents o per cent K 2 0. 
If we take a point half-way between these tw r o points, for instance 
point 61, we have a mixture of the two salts in equal proportions; 
i.e., the fertilizer constituents represented by that point will be 50 per 
cent K 2 and 50 per cent NH 3 . Similarly, point 16 represents 50 
per cent K 2 and 50 per cent P 2 s , and point 21 represents 50 per 
cent NH 3 and 50 per cent P 2 O s . If we take a point nearer to either 
of the corners, w r e will have a higher percentage of that ingredient 
and a correspondingly lower percentage of the other. For instance, 
at the point 59 the composition is 70 per cent K 2 and 30 per cent 
NH 3 ; at 29 it is likewise 70 per cent K 2 0, but 30 per cent P 2 5 ; at 
64 it is 20 per cent K 2 and 80 per cent NH 3 ; at 45 it is likewise 
80 per cent XH 3 , but 20 per cent P 2 5 . 

As stated above, points on the base line 56 to 66 represent mixtures 
containing no P 2 O s . The next line above this, namely 46 to 55, 
represents mixtures containing throughout 10 per cent P 2 O s , but 
varying amounts of the other two constituents. Similarly, the line 
37 to 45 represents throughout 20 per cent mixtures of P 2 5 ; line 
29 to 36, 30 per cent mixtures of P 2 5 ; and so on upward until point 
1, the apex of the triangle, is reached, where the composition is 100 
per cent P 2 5 , as already explained. Similarly, points on the line 1 
to 66 represent o per cent K 2 0; line 2 to 65 represents 10 per cent 
K 2 0, but varying amounts of P 2 O s and NH 3 ; and so on until at 
point 56 the composition is 100 per cent K 2 0. Likewise, points on 
the line 1 to 56 represent o per cent NH 3 ; line 3 to 57 represents 
10 per cent NH 3 , but varying amounts of P 2 5 and K 2 0; and so on 
until at point 66 the composition is 100 per cent XH 3 . It is obvious, 

5 Such diagrams for physical chemical work, giving still finer rulings, namely 
100 to each line, can be purchased from the Cornell Co-operative Society, and were 
used in these investigations. 


therefore, that any point within the triangle represents a mixture 
composed of the three constituents, its position in the triangle being 
determined by the composition of the mixture, namely the ratio of the 
three component parts, P 2 5 , NH 3 , and K 2 0. For instance, point 
12, being on the 60 per cent phosphate line, represents this composition 
of P 2 O s , namely 60 per cent, and being at the same time on the 10 per 
cent NH 3 line, and the 30 per cent K 2 line, it represents 10 per cent 
and 30 per cent of these constituents, respectively. The composition 
of the mixture represented by this point is, therefore, P 2 O s 60 per 
cent, NH 3 10 per cent, K 2 30 per cent; i.e., the ratio of these con- 
stituents in the fertilizer mixture is 60-10-30 or 6-1-3. Similarly, 
the point 34 represents the following mixture of the composition: 
P 2 O s 30 per cent, NH 3 50 per cent, K 2 20 per cent, or a fertilizer 
ratio of 3-5-2. 

For the sake of convenience in stating such ratios or percentage 
composition of the fertilizer mixtures in this investigation, the figures 
are always given in the order P 2 5 , NH 3 , and K 2 0, as shown above. 

The triangle, therefore, represents single fertilizer constituents 
at the apices or vertices, mixtures of any two constituents along the 
boundary lines of the triangle, and mixtures of all three constituents 
within the triangle. 

Solution culture methods 


Before proceeding with a description of the general appearance 
of the cultures growing in solutions with the different fertilizer ratios, 
it may be well to describe briefly here the solution-culture method 
used in these experiments, as the manner of procuring a sufficiently 
large number of seedlings and of preparing physiologically pure water 
are important factors in carrying on this or allied investigations. 


In the work under consideration, as well as in other work in prog- 
ress in these laboratories, it is necessary to have a large number, often 
several hundred and sometimes thousands, of uniform seedlings; 
i.e., seedlings of the same age and equal in development and general 
vitality. The general principle on which this equal germination of 


wheat seedlings is based has already been described by Livingston 6 
and in former bulletins of this bureau. 7 The method consists in 
having a perforated disk, supported by ordinary corks in such a way 
that it will just float upon the surface of a pan of water. In the earlier 
w r ork a wire gauze, w r hich had been coated with paraffin so as to make 
virtually a plate of paraffin reinforced by wire, was used, holes being 
made in this plate by means of a hot wire. Perforated cork sheets, 
preferably paraffined, have also been used. In this case sheet cork 
about one-eighth of an inch in thickness is immersed in melted paraffin, 
and after removal from the paraffin this is allowed to harden in the air 
spaces of the cork. Holes are then made by means of a small cork 
borer so as to give a perforated plate. Both of these plates are open 
to objections. In the paraffined plate, when it is used continually, 
there is a tendency for the paraffin to split off from the wire, which is 
thus exposed to the action of the water and the roots; moreover, this 
plate is not easily repaired. The cork plate, on the other hand, shows 
considerable tendency to warp, is rather fragile, and is not easily 
kept sterile. 

Instead of these plates there were used in these present experiments 
perforated hard-rubber sheets, thus overcoming the above objections 
to a considerable degree. These are prepared by cutting a circular 
disk of 305 mm. from vulcanized sheet rubber 3 . 2 mm. in thickness. 
By clamping several of these tightly together, preferably between 
layers of wood, small holes are drilled through the mass approximately 
4.8mm. in diameter and 5.0mm. apart. Disks of this material 
float level upon the water when supported by corks, are not so sub- 
ject to warping, and are readily cleaned and kept sterile. The corks 
are fastened to the under surface of the disk in four or five places on 
the circumference and in the center. For this purpose either rubber 
or wooden pegs or wire may be used; if the latter, it should be made 
either of iron or aluminum, never of brass or copper. The size of the 
corks is so gauged by trial that the disk is just supported on the surface 

6 Livingston, B. E., A simple method for experimenting with water cultures. 
Plant World 9:13. 1906. 

7 Schreixer, O., and Reed, H. S., Some factors influencing soil fertility. Bulle- 
tin 40, Bureau of Soils, U.S. Dept. Agric. 1907. 


of the water. After floating this perforated disk on the surface of the 
water, the wheat seed, which has been previously soaked for about 


two hours, not longer, is spread evenly over the surface. This method 
insures an even germination of the seed and has the advantage of 
keeping the seed and later the young seedlings just moist. The seeds 
are never submerged in the w r ater, nor do they remain suspended 
dry above the water. The roots grow through -the holes of the disk 
into the liquid below. This method of sprouting wheat seedlings is 
far superior to growing them in sand, since the seedlings are more 
uniform and the apparatus can be kept sterile much more readily 
than sand. Furthermore, for water culture purposes the seedling 

y lifting it from the disk. In this way 



removing seedlings trom sand some injury w r ould be unavoidable. 
Moreover, the seedlings are removed direct from water to grow in 
solutions, and thus have the advantage of germinating in a medium 
similar to that in which they are to be grown. 

The water in the germinating pans may be distilled water, if 
desired, or where good tap water can be secured this may be used. 
The water in the pans is changed daily, or, during the warm weather, 
twice a day. Several hundred uniform seedlings can be procured 
from a single disk of the kind described. Whatever inferior plants 
occur are rejected. For the bottle culture work described in this 
bulletin, the seedlings were used when the plumule was about 2 cm. 
high and just ready to emerge" from the enveloping sheet. 

More recently disks of perforated aluminum have been used in 
this laboratory and have proven very satisfactory. These disks are 
floated by means of a raft of sealed glass tubing of such dimensions 
as is required to float the plates in the manner above described. 
Three lengths of tubing 102 cm. long, 35 mm. in diameter, with 
approximately a 1 . 5 mm. wall thickness, were sufficient to float six 
aluminum disks 30 cm. in diameter and 1 .6 mm. in thickness. The 
lengths of sealed tubing and two glass rods 56 cm. long and 12 mm. 
in diameter are wired together into a raft. The entire arrangement 

is floated in a porcelain-lined iron tank. Fig. 2 shows this tank with 
seedlings at various stages of development. 





The bottles used in these cultures are made of flint glass, have a 
capacity of about 250 cc, a total height of 100 mm., an outside diam- 
eter of 70 mm. at the bottom, a neck about 20 mm. high, and a mouth 
57 mm. in diameter. This bottle is stoppered by means of a soft, 
flat cork about 12 mm. in thickness and notched for holding the 
seedlings. The method of notching these corks consists in cutting 
ten vertical, triangular wedges from the circumference. Each wedge 
after being cut out is truncated, so that when replaced a small tri- 
angular opening, through which the plumule of the seedling will pass, 

Fig. 2. — Method of germinating wheat seedlings on aluminum disks, which are 
floated in an enameled tank; disk to right elevated to show the plant roots. 

is formed. This hole should be large enough to hold the seedling 
firmly and yet not bruise or injure it in any way by pressure. Around 
the circumference of the cork and in the upper half a groove is made 
sufficiently large to hold a small rubber band. After the wedges are 
inserted, the band keeps them in place and allows the cork with the 
seedlings to be handled readily and put into or taken out of the bottle 
without disturbing the plants. The seedlings are most easily inserted 
in the cork in the following manner: The cork with the ten wedges 
held in place by the rubber band is taken in the left hand and the seed- 
ling in the right hand. The plumule is pushed through the small, 
triangular opening, and then pulled up until the seed is close against 
the cork. When older seedlings, in which the leaves have unfolded 


themselves, are to be used for an experiment, or where plants such as 
cowpeas are used, the wedge must be removed, the seedling put in 

place, and the wedge and rubber band replaced. 


It is safe to say that ordinary distilled water, such as is commonly 
found in laboratories, is unsuited for culture experiments. This is 
due to a variety of causes which need not be entered into here, but 
which are discussed in a bulletin by Livingston. 8 Such water, while 

pure from a chemical and physical standpoint, is nevertheless not 
suited for such delicate indicators as plants, which are highly sus- 
ceptible even to very minute quantities of toxic materials. Such 
water can be improved by distillation with strong oxidizing agents, 
such as potassium permanganate in alkaline solution or potassium 
bichromate in acid solution. The far simpler method already 
described by Livingston 8 has given very satisfactory results in this 
laboratory. In this method the water is purified by shaking it with 
a highly absorptive carbon black, which removes from the water any 
traces of injurious bodies it may contain. Not all carbon black 
possesses this property to an equal degree; the variety used in this 
laboratory is known as the "G Elf" brand, and is prepared on a 
commercial scale by burning natural gas and condensing the finely 
divided carbon on cooled surfaces. It is the same carbon black used 
for the purpose of decolorizing soil and plant solutions in this labora- 
tory. For the present purpose the carbon black is thoroughly washed 


removed from it or not. The 


so as to form a thin paste. About 10 cc. of this paste is added to each 
liter of water which is to be purified, and after shaking the mixture is 





satisfactory for the grow 

8 Livingston, B. E., Further studies on the properties of unproductive soils. 
Bulletin 36, Bureau of Soils, U.S. Dept. Agric. 1907. 




General behavior of the cultures containing different 

fertilizer ratios 

The plants in the different cultures prepared as described in the 
previous section show some marked differences almost from the 
very outset. After several changes of solution have been made, these 
differences become quite prominent. The most marked of these is 

P Z°5 


Fig. 3. — Green weight of 10 wheat plants grown in 66 cultures with different 
ratios of P 2 5 , NHj, and K 2 (). 

that the solutions containing only one or two of the fertilizer ingredients, 
i.e., the entire periphery of the triangle, show a markedly less develop- 
ment of the plants than the cultures in the interior of the triangle 
wherein the three elements are combined. In other words, even the 
addition of only 10 per cent of the third element to mixtures of the 
other two produces a very appreciable increase in plant growth. 
This effect is most noticeable in the change from no nitrate to 10 per 


cent nitrate. Of the single fertilizer ingredients, the nitrate culture 
is the best, and this is usually followed by the potassium culture, the 
phosphate culture being in general the poorest of the entire triangle. 
The cultures in the interior of the triangle do not show so marked a 
difference, although it is apparent that there is a tendency toward a 
rise in the middle of any line, this being displaced somewhat along 
some of the lines so as to give the effect of the highest and better region 
of growth lying in the middle of the lower part of the triangle; i.e., 
nearer to the potassium-nitrogen base. The set grew from February 
25 to March 21. The green weights obtained in the sixty-six cultures 
are given in the triangular diagram shown in fig. 3. A more detailed 
discussion as well as grouping of these green weights, their correlation 
with the concentration of the solution, and the amount of nutrients 
removed will be given later. 


As has already been stated, the solutions were analyzed every 
third day for the three component fertilizer parts, phosphate, nitrate, 
and potash, expressed as P 2 5 , NH 3 , and K 2 0. The original con- 
centrations in these elements were in the sum total 80 parts per mil- 
lion. After the analysis the sum total of the three component parts 
was again calculated and the average concentrations of these three 
elements was again ascertained for the eight periods. The average 
concentrations will be found in the diagram in fig. 4. It is thus 
apparent that more of the essential ingredients were removed in the 
interior of the triangle; i.e., there was a greater absorption where 
all three were combined than where only two elements were present, 
and the least removal took place in those cultures where the single 
salts were present. It is also noticeable that in the solutions contain- 
ing the three fertilizer elements the greatest removal occurred in that 
region of the diagram where the greatest growth occurred. A more 

detailed discussion of these results and their component parts will 
be given later in this paper. 

Ratios of P 2 5 , NH 3 , and K 2 0, found in the various cultures 

It might be said that in all cases the ratio in the final solution was 
never the same as the original ratio. The amount of change which 
had taken place, however, was markedly different in the different 
solutions and depended largely upon what the original ratio was. In 




order to show this change in ratio of the fertilizer ingredients, the 
triangular diagram is of the greatest service, for without this it would 
be impossible to get an intelligent idea of what had occurred in the 
solution. In the diagram fig. 5 are given the original ratios of the 
fertilizer constituents, the ratios left in these solutions as shown by 
analysis, and the corresponding ratio of the removed constituents. 


Fig. 4. — Showing the average concentration in parts per million of P 2 O s , NH 3 , 
and K 2 of the solution after the growth of 10 wheat plants; concentration of the 
original solution was 80 ppm. 

It may be shown mathematically that these three points lie upon a 
straight line and this is the case in the diagram. 

The large dots in the diagram represent the original ratios corre- 
sponding to the scheme previously explained and given in fig. 1. The 
circles indicate the ratio left in the solution as shown by analysis. The 

.f th- 

ing ratio of the removed materials. 

It must be borne in mind that 



this diagram deals only with the ratios of the ingredients and not with 
the amounts that are present. 

It is apparent that there is a decided tendency for these lines to 
converge toward a region somewhat below the center, as shown by 


the congregation of arrow 7 points in this region. In other words, the 
solutions near this central area change least in their ratio, and the 

Fig. 5. — Showing the ratio of the original, the final, and the ratio of the loss of 
P 2 O s , NH 3 , and K 2 from the culture solution: the dots indicate the ratio of the 
constituents in the original solution; the circles show the ratio of the constituents in 
the solution after growth; and the arrows show the ratio of the decrease. 

farther the ratios were removed from this central area originally, the 
more were they altered in the course of the experiment According 
to the diagram this area would seem to lie between the 10 and 20 per 
cent phosphate line, since the points on the lower line have moved 
in a direction opposite to that taken by the ratios in the upper part 
of the triangle. The data already presented show in general that 
the area of greatest growth occurred in this same region. It is in 


this region of greatest growth, therefore, that the greater absorption 
took place with the least change in ratio; in other words, the solutions 
represented by this region offered the best environment for plant 
development and the best ratios for the absorption of plant nutrients. 
Attention might also be called to the fact that in the ratios where 
the nitrate was low, there has been a movement to the no nitrate line, 
or at least so close to this line that it was impossible to plot them other- 
wise. After the comparatively small amount of nitrate was removed 
or reduced to a minimum, the point marking the ratio would have 
to move along the no nitrate base line in the direction either of the 
potash or of the phosphate, depending upon which was removed in 
the larger amount. The diagram shows that this movement of the 
ratio in the solutions was in nearly all cases toward the phosphate 
apex of the triangle. It is obvious that such a condition of affairs 
will cause a shifting of the lines connecting the ratios in the diagram, 
and that a similar state of affairs in the cultures on the low potassium 
line would produce a shifting of the ratio lines in the opposite direction. 
An examination of the diagram shows this has occurred, because the 
ratio lines in the upper part of the triangle show a divergence, giving 
a fanlike effect. In the case of the potash this divergence is also 
strongly noticeable, although the potash was never so completely 
removed as was the case with the nitrate. 

The results of the experiments considered by periods 

The very earliest periods, when compared with the later periods, show 
some differences in the ratio in which the three elements are removed, 
as has already been pointed out. Mention has been made of the fact 
that the ratios removed from the solutions along the lower phosphate 
line, namely, the 8 parts per million line of cultures, had a tendency 
to cause the ratio lines to turn from a point below this line to points 
above the line with increase in time. This general behavior of the 
cultures is very well shown by the three diagrams for the first, second, 
and third periods. The first diagram (fig. 6) shows the arrow points 
of the io per cent phosphate mixtures to lie below this line. The 
second diagram (fig. 7) shows some of the points above and some 
still below the line. In the third diagram (fig. 8) all but one arrow 
point lie above the line, and some arrow points of the 20 per cent 



phosphate line are also noticed to have turned so far that some of 
them take a position even slightly above this line. There is a general 
tendency, moreover, for all of the points to lie somewhat higher for 
the other cultures, thus showing that this influence is probably com- 
mon to all the cultures, though seen most strikingly in those contain- 


small amounts, where an actual reversal takes place. 



Fig. 6. — Showing the ratio of the original, the final, and the ratio of the loss of 
P 2 O s , NH 3j and K 2 from the culture solution in the first period. 

general result may be interpreted to mean that the relative amount 
of phosphate removed in the early stages of the seedlings is not so 
great as it is subsequently, and the phenomenon can probably be 
correlated with the fact that in experiments where distilled water is 
used rather than these solution cultures, phosphate is excreted by 
the germinating seed and can be detected in the water in which 
germinating seeds or young seedlings bathe, 
when used in these experiments have passed in the main through this 

Although the seedlings 




stage, the results show that there was a material lessening of the power 
to remove phosphate, probably due to the fact that the process of the 
phosphate absorption had not fully replaced the opposite function 
existing during germination and early growth. 

The potassium absorption is also different from that in the later 
periods, although this is not so striking in the diagrams as in the case 



Fig. 7. — Showing the ratio of the original, the final, and the ratio of the loss of 
P 2 O s , NH 3 , and K 2 from the culture solution of the second period. 

of the phosphate. The average ratio of the removed material during 
these first three periods for all the cultures is 19 for phosphate, 39 for 
nitrate, and 42 for potassium; whereas the average for the five succeed- 
ing periods is 21 for phosphate, 44 for nitrate, and 35 for potash, thus 
showing that there is a tendency for a relatively greater potash and 
a relatively less phosphate absorption to take place during the earlier 
periods. So far as the influence of these fertilizer elements on growth 
is concerned, however, it is to be noticed that the nitrate has the 



greatest effect, as is shown by the fact that the difference between 
the no nitrogen line and the 8 ppm. line is very great in the case of 
the nitrate, less in the case of the potash, and least in the case of the 
phosphate. As a rule, beyond the second or perhaps the third period 
the diagrammatic representation of the result is on the whole uniform, 
but is influenced undoubtedly by the conditions of growth during any 


Fig. 8. — Showing the ratio of the original, the final, and the ratio of the loss of 
P 2 5 , XH 3 , and K 2 from the culture solution in the third period. 

period; in other words, by weather and other conditions, which is 

removal from 


ence of bacteria. 

\ course of these experiments bacteria and 
other microorganisms were excluded so far as possible, but no special 
effort was made to maintain absolutely sterile conditions, inasmuch 
as this would have been a practical impossibility in an experiment on 
so large a scale. Moreover, it may even appear questionable whether 


absolute sterility would be desirable. The bottles were sterilized 
before being used in making culture solutions for the various changes, 
the pans and other apparatus used in germinating the seed were ster- 
ilized from time to time, and corks used for the cultures were always 
clean and sterilized before use. Although all of these precautions 
were taken, it was of course not possible to exclude some micro- 
organisms in such work, as the solutions were exposed from time to 
time to the air. There was in no case any excessive microorganic 
life noticeable. While bacteria and other microorganisms were 


present in the cultures to a slight and, under the conditions, unavoid- 
able extent, it can hardly be said that their influence could have been 
large; that is, such influence as they had was probably so slight as 
to be negligible so far as the general and larger tendencies which are 
shown in this paper to exist are concerned. 

Controls which were set up without having plants grown in them 
were found not to have changed their concentration or proportions 
of the constituents. Moreover, the various cultures support each 
other in their general tendencies. If the disappearance of nitrates 
were to be ascribed to bacterial activity, this should have shown itself 
all the more prominently as the age of the cultures increased. Such 
changes as were noticed from period to period might be ascribed to 
changes in the climatic conditions, thus still further affecting 
the plant's metabolism. This is very nicely illustrated in a series 
of preliminary experiments in which the solutions were changed 

every day instead of every 

The dia- 

gram giving the results obtained on a clear day shows strikingly a 
relatively greater nitrogen removal under these conditions. The 
diagram giving the results obtained on the following day, which was 
cloudy and rainy, shows no less strikingly that relatively less nitrogen 
was absorbed on the cloudy and rainy day. It would be obviously 
unfair to conclude that under such conditions bacterial activities 
had been greater on the sunny day than on the following rainy day, 
especially as this result is in harmony with all observations on the 
removal of nitrate from solutions by plants. The processes of nitrogen 
utilization within the plant are known to be greater under conditions 
of better illumination. It is fair to assume, therefore, that this same 
general tendency held in all the other cultures, and that whatever 



effect the bacteria may have had, it was slight in comparison to the 
activities of the plant itself under the conditions of this experiment. 

growth and concentration of P 2 5 , NH 


The experimental results reported seem worthy of a closer analysis 
than that accorded them thus far. General tendencies as indicated 

Fig. 9. — The arrangement of the culture solutions in groups: a, b, c, three 
fertilizer element groups; d, two fertilizer element group; e, one fertilizer element 

by the individual cultures have already been pointed out, and these 
general impressions gleaned from the individual results can be better 
shown by a method of grouping. This method of grouping should 
be particularly valuable in showing the general tendencies, either in 
regard to plant growth or concentration differences connected there- 
with. Such a system of grouping, comparing different regions of the 
diagram, can be made in various ways. 




The grouping shown in fig. 9 was made because by so doing the 
area of the greatest growth, which was always somewhat below the 
center of the triangle, will fall in these experiments into the central 
group. This central grouping is shown in the diagram as group 0, 
the connecting lines showing the cultures included in this group. 
Group b takes in the cultures lying immediately outside of this cen- 
tral group. The same applies to group c y which is, however, farther 
removed from the central group. It will be noticed that these three 
groups contain the three fertilizer constituents. The next or fourth 
group contains all of the cultures in which two fertilizer elements occur, 
as shown by the lines. The fifth and last, group consists of the sum 
of the single fertilizer elements. In handling the date, therefore, the 
average green weight in grams for each of these groups was deter- 
mined, together with the average final concentration in the respective 
fertilizer constituents, or their combinations. In table I the average 
green weight and combined concentration of P 2 5 , NH 3 , and K 2 
is shown. 

Average green weight axd concentration of P 2 5 +NH 3 + K 2 obtained in 


(ppm. = parts per million) 

Culture solutions 

Three fertilizer elements, group a . . 
Three fertilizer elements, group b. . 
Three fertilizer elements, group c. . 

Two fertilizer elements 

One fertilizer element 

Three fertilizer elements, groups 


All cultures combined 













2 .155 
1 . 502 








Average P 2 O s . NH* 
and K 2 




29 .O 



68. S 






5 6 -3 




43- 2 

In the third column the relative green weight for the different 
groups is given, taking the results in group a as ioo. From this it 
will be seen that the growth in group b is 88 per cent of that in group tf, 
group c is 75 per cent; in the two fertilizer element group it is 46 per 
cent, and with the single fertilizer element the average result is only 



34 per cent. The concentrations in these various groups are shown 
in the fifth column; the original concentration in all these cases was 
80 parts per million of the combined constituents. In the first group, 
which was the one giving the greatest growth, the concentration was 
decreased from 80 to 29 parts per million, although, as has already 
been shown, the ratio of the three constituents in the solutions of this 
group suffered less change than did the ratio in the other groups. 
This can be seen from the diagram in fig. 5. In the second group the 
concentration was reduced to 35 parts per million; in the third group 
of the three fertilizer constituents to 44 parts; in the group of two 
fertilizer constituents it was reduced to 54 parts per million; and in the 
group containing one fertilizer constituent it was reduced to only 
69 parts per million as an average. 

For the sake of comparison, a group containing all cultures having 
the three fertilizer elements, as well as the combination of the entire 
group of sixty-six cultures, is given. The table also very clearly 
shows the markedly different effects produced on the green weight 
by a single fertilizer element and when these are used in combinations 
of two and of three, the results being approximately 1.5, 2.2, and 
4.0, respectively, the higher group within the latter being as high as 

It will also be of interest to consider this same grouping in respect 

to the concentrations of the fertilizer elements individually. The 

number of cultures included in the group where two fertilizer elements 

were used was necessarily reduced, inasmuch as the ingredient in 

question did not occur in nine of the solutions, and the group of single 

fertilizer elements becomes, of course, reduced to only one culture, 

and for this reason has very little comparative value, although the 

result is usually decided enough to allow no doubt as to the general 

In the fifth column of table II is given the concentration of 
found after growth, the original concentration being given 
in the fourth column for comparison. In the last column is shown 
the percentage decrease in the concentration of P a O r This per- 
centage decrease was in the first group 50 per cent, in the second 
group 37 per cent, and in the third group 27 per cent. In the combi- 
nation of two fertilizer elements, one of which was phosphate, it 

P 2 O s 




was 17.5 per cent, and in the phosphate by itself it was only 10 per 
cent. The actual decrease in concentration appears from the figures, 
however, to be strikingly uniform, varying in the different groups 
from 7 to 9 parts per million. This relationship of final to original 
concentration, however, is not properly shown by this form of grouping 

Average green weight and concentration of P 2 5 obtained in different 


Culture solutions 

Three fertilizer elements, group a 
Three fertilizer elements, group b 
Three fertilizer elements, group c 

Two fertilizer elements 

One fertilizer element 

Three fertilizer elements, groups 










4 645 


1 -S3 1 








Average P a 5 


26 .0 




7 i.8 









and can be much more satisfactorily handled, as will be done later, 
by grouping only such phosphate solutions as have the same original 
concentration. In the present table the original concentration is 
really an average of points varying, for instance, in the third group 
from 8 to 64 parts per million in phosphate content, giving the average 
concentration of 34 parts per million shown in the table. 

Average green weight and concentration of NH 3 obtained in different 


Culture solution- 

Three fertilizer elements, group a 
Three fertilizer elements, group b 
Three fertilizer elements, group c 

Two fertilizer elements 

One fertilizer element 

Three fertilizer elements, groups 











4 • 645 




















2.75 2 






• • 







43- 1 

66 5 



In table III is given the same character of results as have been given 
in the preceding section for phosphate, except that the concentration 
of nitrates is considered. 

In the groups of two and single fertilizer elements, only those cul- 
tures containing nitrate are considered. In the three groups com- 
prising the three fertilizer ingredients, the percentage decrease in 
nitrate is seen to be respectively 69, 70, and 60; in the two fertilizer 
element group it is 43; and in the single element, 12. 

In table IV are given the results for the potash concentrations, 
similarly arranged and grouped as in the preceding tables. 

Average green weight and concentration of K 2 obtained in different 


Culture solutions 

Three fertilizer elements, group a 
Three fertilizer elements, group b 
Three fertilizer elements, group c 

Two fertilizer elements 

One fertilizer element 

Three fertilizer elements, groups 








in grams 






1 .106 



Average K a O 







• • 

22 .9 












The percentage decrease in the cultures is for the three fertilizer 
elements in the three groups 66, 58, and 58, respectively; for the 
two fertilizer elements, 35; and for the single fertilizer element, 45. 

Influence of different amounts of P 2 5 , NH 3 , and K 2 0, varying 

respectively, from 8 to 80 parts per million 

In the consideration of the results up to this point, no special dis 
cussion has been made in regard to the actual amounts of the individ- 
ual fertilizer ingredients which were in the cultures. By reference 
to the original description of the diagram fig. 1 it will be seen, for 
instance, that there was a series of cultures in the line from number 
46 to number 55 which contained 10 per cent of phosphate in the 
fertilizer mixture, or 8 parts per million of the culture solutions. Simi- 
larly, the line 37 to 45 contained 16 ppm.; the line 29 to 36, 24 ppm.; 




and so on to the apex of the triangle. In the same manner, the line 
of cultures from 3 to 57 contained 8 ppm. NH 3 as nitrate; the line 

6 to 58, 16 ppm.; and so on up to 80 ppm. NH 3 in culture 66. 


line of cultures from 2 to 65 contained 8 ppm. K 2 0; line 4 to 64, 16 
ppm.; and so on up to 80 ppm. K 2 in culture 56. 

It will be interesting to see what effect these different amounts 
of each fertilizer ingredient have had on the growth of the plant, and 
the removal of this constituent during the period of growth. In such 


Showing the results obtained for P 2 O s in tele cultures containing different 

amounts of p2o5 (ppm. = parts per million) 

of P 3 O s 










Parts per 

P 3 O s 


content of 

P,0 5 




Average decrease of 

p 2 o 5 












61 -5 



1 .226 

• 64 




J 7-5 






3 • 045 






3 2 



22 .2 








3- 2 95 


* * • • 

* * • 

• • ■ • 

3 135 

P,O s 
decrease per 







3- 2 



• » 




in the cultures along any of these lines has been studied and is pre- 
sented in this section. With this in view all the results obtained in the 
various cultures in any one line were added together, and as the 
number of cultures naturally differ from line to line, the average 



is not intended to mean that all the cultures along this line were simi- 
lar, for it has already been showp that in general the points in the 
middle were higher than those at the end of any one of these lines. 
Phosphate. — In table V are given the results obtained for P 2 
in the cultures containing different amounts of P 2 5 , varying fron 

80 to 8 DDm., as shown in the second column of the table 



The first column gives the percentage composition of the fertilizer 
mixture so far as the P 2 O s content was concerned. In the third 
column is given the average concentration for the eight three-day 
periods in the experiment, and in the fourth column, the difference 
between this and the original concentration, thus giving the average 
decrease, which is expressed as a percentage in the fifth column on 
the basis of the original concentration. The sixth column gives the 
average green weight obtained along any of these concentration lines, 
and the next column gives the decrease in P 2 O s calculated to the unit 
basis of one gram production of green plant. In other words, the 
decrease recorded in the fourth column has been divided by the 
green weight corresponding thereto in the sixth column. This gives, 
as it were, the rate of decrease in parts per million of the solution for 
each gram of green weight produced; but if it is desired to have the 
result expressed in terms of milligrams of P 2 O s removed by each 
gram of green weight produced, the figures in the column must be 
multiplied by 2, since 250 cc. of solution were presented to the plants 
eight times; i.e., a total of 2000 cc. It will be noticed that the green 
wxight steadily increased as the phosphate content decreased. It 
must be borne in mind, of course, that when the phosphate content 
decreases, there is a corresponding increase in the average content 
of both potash and nitrogen. In the last 'figure for the green weight, 
however, namely that in which the phosphate content became zero 
in the culture solutions, there is again a marked drop, although the 
potash and nitrogen content in these was higher than in any of the 
solutions above this in the table. It follows accordingly that a very 
distinct part was played by the phosphate in producing growth, 
although its maximum efficiency seems to be reached in these experi- 
ments in rather low concentration. Attention might also be called 
here to the fact that the concentrations of phosphate in soil solutions 
are always low, and relatively much lower than any of the other 
constituents here considered. The plant in its natural environment, 
therefore, has adapted itself to the occurrence of this constituent in 
weak solutions. 

The increase in green weight shown by the table to be from 3 . 1 
to 3.6 grams corresponded to an increase from o to 8 ppm. in the 
original P 2 O s content. The further increase to 16 ppm. P 2 O s has 


produced no further increase in plant growth, and above this a decrease 
has even followed, which may be in part due to some direct influence 
of the phosphate solution, the carrier here used being calcium acid 
phosphate, and may also be due in part to the fact that the quan- 
tities of potash and nitrate are being decreased as the phosphate 
increases. This relation, however, has been discussed in a much 
more thorough manner where the ratios of these substances are con- 
sidered, and the change in concentration which the solutions undergo, 
as shown in the diagram of ratio change in fig. 5, makes this matter 
very clear. It is there shown that the region of better growth is 
found in the lower part of the diagram, and this is also the region 
of most absorption and least change in the ratio. All these regions 
it will be noticed comprise solutions which contain low amounts 
of P 2 5 . 

The average concentration of phosphate in the culture solution 
after growth, as shown in the third column, continually decreases. 
The actual decrease in the next column, however, shows that the 
greatest decrease in concentration of this element does not go with 
the greatest growth, but is found in the 40 ppm. phosphate solu- 
tions, increasing steadily from the 8 ppm. solution to this point, 
and then becoming practically constant up to the point where the 
other fertilizer elements entirely disappear and only phosphate 
remains, when it again drops slightly, the conditions for growth also 
being much poorer through this total absence of potash and nitrogen. 
The decrease when expressed as a percentage of the amount of mate- 
rial originally present shows that the removal is the more complete 
as the original concentration is lower. The last column in this section 
of the table, giving the decrease per unit of green weight, shows that 
equal weights of green plants cause quite unequal decreases in con- 
centration, this change being least in the weaker solutions and greatest 
in the higher, thus indicating that more was removed by the plant 
in the higher concentrations than it could economically utilize under 
the conditions; i.e., the plant absorbed the material because it was 
there, although it apparently had already all that it could utilize 
economically. This is a fact consistent with field observations and 
ash analysis, where it is frequently noticed that the actual amount 
of mineral constituents is larger in the poorer plants. 



Nitrate. — Table VI gives the results obtained for nitrate in the 
cultures containing different amounts of nitrate, similarly arranged 
and computed as in the phosphate table just discussed. 

Showing the results obtained for NH, in the cultures containing different 



of NH 3 

Parts per 
million of 

NH 3 



content of 

NH 3 




Average decrease of 

NH 3 



NH 3 























42 .O 
12 .O 



• 0.2 

m # * « 


22 .O 


21 .7 



r • * • 







* • 1 



• 4 

















3- 6 • 







• • • 

The green weight column differs from the one discussed in the 


phosphate table in that the highest result occurs with the 32 parts 
per million NH 3 line, the numbers descending in each direction from 
his higher green weight. The change from o to 8 ppm. NH 3 is very 
marked in the green weight, which increases from approximately 
1 . o gram to 3 . 2 grams, an even greater change than was noted in 
the phosphate table. The average decrease in concentration w r as in 
this case greatest between the limits of 32 to 64 ppm., the increase 
being gradual from the 8 ppm. solution up to the 32 ppm. solution, 
then being practically uniform up to 64 ppm., when it again declines. 
The figures in the percentage decrease column are on the whole much 
greater than those in the corresponding column in the phosphate 
table, thus showing a comparatively greater decrease of the nitrogen 
than of the phosphate in solutions of equal content of these elements, 
respectively. The last column, showing the rate of decrease per 
unit of green weight, shows the same general tendency for these fig- 
ures to decrease with the decline in the original content of nitrogen 
in the solutions, as was noticed in the phosphate table, except that 




the first two results, namely in the 80 and 72 ppm. solutions, are in 
this case lower than the following three or four entries, while in the 
phosphate table these two were the highest in the column. The 
conditions for growth with the phosphate in these two solutions, or 
groups of solutions, were very poor as compared with the rest of the 
results, while with the nitrate this difference in growth was not so 


Potassium. — In table VII are given the results obtained for potas- 
sium, the table being arranged and the results computed in the same 
manner as with those in preceding tables. 


Showing the results obtained for K 2 in the cultures containing different 




Parts per 


Average decrease of 

K 2 

of K a O 



K a O 




K a O 


K a O TN 









TION (ppm.) 








19 -5 

1 .106 

14. I 


- 54.1 ^ 

[ 17-9 



1 .922 








2 .66l 




3 6 -9 

19. 1 



2 .890 





19. 1 



3 .222 









3 .600 



3 2 


















12 .1 







3- 1 







* • * • *• + # 

« * 

• • 

1 -97^ • ■ * 

The green weight column is in its general tendency very similar 
to that observed in the nitrate table, namely that the 32 ppm. cultures 
give the highest green weight. The figures representing the average 
decreases and percentage decreases, on the other hand, correspond 
in trend more closely with those shown in the phosphate ^able, this 
similarity being especially shown in the last column giving the rate 
of decrease per unit of green weight, in which a regularly descending 
column of figures is seen. In the case of the green weight, the change 
from the 8 ppm. solutions to those containing no potash whatever 
was, as in the other cases, very marked, and this drop in green weight 
was also distinctly noticeable in the solution where potash alone was 
present. The results given in the three columns in the tables showing 


the rate of decrease per unit of green weight are consistent in showing 
that relatively more was absorbed when more w r as present, although 
the plant does not seem to have been able to utilize this increase 
economically in its growth. This tendency is especially marked in 
the case of phosphate and potash, although it is also shown to an 
appreciable extent in the case of nitrogen. 

As w T ill appear in future publications, this general method of 
experimentation was used for the purpose of studying the effect of 
individual soil contituents and other organic compounds by using 
them in uniform concentration in all the cultures of a triangle. In 
these studies it was necessary to grow a control set without the con- 
stituent to be studied, so that the foregoing experiment was in this 
manner repeated a number of times, and the general results thus 
obtained were in harmony with those here recorded. 


In this study the growth relationships and concentration differences 
were observed between solution cultures in which the phosphate, 
nitrate, and potash varied from single constituents to mixtures of 
two and three in all possible ratios in 10 per cent stages. 

The better growth occurred when all these nutrient elements were 
present, and was best in those mixtures which contained between 
10 and 30 per cent phosphate, between 30 and 60 per cent nitrate, 
and between 30 and 60 per cent potash. The growth in the solutions 
containing all three constituents was much greater than in solutions 
containing two constituents, the solutions containing the single con- 
stituent giving the least growth. 

The concentration differences noticed in the solutions were also 
very striking, the greater reduction in concentration occurring where 
the greatest growth occurred. 

The change in the ratios of the solutions and the ratios of the 
materials that were removed from the solutions showed that where 
the greatest growth occurred, as above outlined, the solutions suffered 
the least change in ratio, although the greatest change in concentra- 
tion occurred. 

The more the ratios in these solutions differed from the ratios in 
which the greatest growth occurred, the more were the solutions altered 


in the course of the experiment, the tendency in all cases seeming 
to be for the plant to remove from any and all of these solutions the 
material in the ratio which normally existed where greatest growth 
occurred. This did not actually occur in all cases, owing to the 
unbalanced condition of some of the solutions. 

The results show that the higher the amount of any one constituent 
present in the solution, the more does the culture growing in that 
solution take up of this constituent, although it does not seem able 
to use this additional amount economically. 

In the very early periods the ratio of phosphate absorption is low 
and the potash absorption high, although in final growth the greater 
response is obtained with nitrate, indicating relatively low phosphate 
requirement and high potash requirement of the seedling plant. 

Bureau of Soils 
U.S. Department of Agriculture 

Washington, D.C. 



W. T. Saxtox 

(with plates i-iii and three figures) 

In a previous communication (10) the writer has pointed out cer- 
tain peculiarities in this genus, and in the genus Callitris, which mark 
them off sharply from all other Cupressineae which have been inves- 
tigated. Many phases of the life history have now been fully inves- 
tigated in both genera, and in the main confirm the results previously 
published, although certain points require modification or correction. 
The investigation of both genera has been carried on simultaneously, 
the original intention having been to present the results in one paper; 
it has however been found that the differences between the two are 
more radical than was anticipated, and the present communication 
relates solely to Widdringtonia. A similar account of Callitris will 
be published very shortly (n), in which the results of both investi- 
gations will be discussed. The methods used have not differed 
materially from those previously employed. 

i. The male cone 

The sporophylls are arranged, like the leaves, in decussate pairs. 
Each sporophyll is peltate and bears four microsporangia 1 on the 
proximal side of its stalk. The output of spores from a sporangium 
is about 500. The structure of the sporophyll in a transverse section 
through the insertion of the sporangia is shown in fig. 1. The struc- 
ture of the cone in radial longitudinal sections has already been 
figured (10, fig. 2). The anatomy of the sporophyll agrees exactly 
with that of Callitris and will be later described in that genus. 


2. The female cone and the ovule 

The young female cone consists of two spreading decussate pairs 
of sporophylls, each scale bearing seven or eight ovules on the proximal 
half of the UDDer surface, and close to the median ridse. 2 Each 

1 Masters (6) says 2-3. 

2 In W. Schwarzii only three ovules are borne on each scale. 


[Botanical Gazette, vol. 50 


megasporophyll is traversed by a large number of vascular strands 
and resin canals; near the top of the sporophyll the latter are evenly 
disposed all round the periphery, and inside each canal is a vascular 
bundle with the phloem on the side nearest the canal. On the lower 
(outer) surface of each scale is a small umbo. 

The structure of the ovule before and shortly after pollination has 
already been described and figured 3 (10, figs, i and 3). The ingrowth 
of the micropyle-closing cells is not followed by septation in them, 
as it is for example in Pinus. A transverse section of the micropyle 
is shown in fig. 2, in which it may be seen that the opening is often 
not entirely closed by the ingrowth of these cells. This figure shows 
a most remarkable resemblance to a similar section of the micropyle 
of Bennettites, figured by Lignier (6, fig. 31), as pointed out to me 
by Professor Seward. 

Some time after pollination the sporogenous tissue becomes evident. 
Seen in median section it appears as about 18 or 19 large cells with 
dense cytoplasm and large lightly staining nuclei, sharply marked off 
from the surrounding cells, which have vacuolate cytoplasm and 
smaller deeply staining nuclei. The position and structure of these 
cells are indicated in figs. 3 and 4. Their number 4 is about 64, 
possibly exactly that number, in which case they are not improbably 
formed by repeated simultaneous divisions of a single archesporial 
cell. It is however quite as likely that the number is somewhat less 
than 64, and that they have not a common origin from a single 
archesporial cell. They are all exactly alike when first differentiated, 
and their subsequent behavior indicates that they are spore mother 
cells, only one of which, however, is functional. Sometimes a single 
layer of tapetal cells may be seen a little later. 

At a somewhat later stage, one of the central mother cells becomes 
a very little larger than the remainder, the nucleus also enlarges very 
slightly, and the wall becomes a trifle thicker than that of its neighbors. 
This is the functional megaspore mother cell. At about the same time 
the non-functional cells elongate somewhat, and certain peculiar 

3 The wings of the ovule were accidentally described (p. 166) as being only one 
cell thick. This is never the case, even close to the margin. 

4 Calculated on the assumption that the actual number is to the number seen in 
median section as §7rr3 to irr 2 . 


structures make their appearance at or near their ends. These struc- 
tures appear as homogeneous dark brown bodies of various shapes, 
and may be conveniently called the "brown bodies" (fig. 5). That 
they are not produced artificially is clearly indicated by the fact that 
they are found only in the sporogenous tissue, and at first only in a 
very definite position at the two poles of the cells. Later they may 
be somewhat displaced from this position. 

The only structures previously recorded as occurring in megaspore 
mother cells, with which the " brown bodies " might conceivably be cor- 
related, are the "kinoplasmic centers" found in Ginkgo (Carothers 
1) and other gymnosperms. It seems unlikely, however, that the 
"brown bodies" are of this nature, especially as their existence has 
not been proved in functional mother cells. They seem to be 
entirely unstained by Delafield's hematoxylin, and show a marked 
resemblance in this respect, as well as in color and consistency, to 
certain nucleoli, although nucleoli usually stain deeply with this 
reagent. At about this time some of the cells in or near the central 
axis of the nucellus begin to break down or to elongate, no doubt 
in preparation for the rapid growth of the pollen tube and embryo 

sac (fig. 6). 

3. Megasporogenesis 

About a week after the events described have taken place the 
functional megaspore mother cell is found in synapsis (figs. 7 and 9), 
the surrounding cells retaining more or less their former character. 
No preparation has been obtained which shows either of the reducing 
divisions. Figs. 10 and 11 show a state of things rather difficult to 
interpret. The arrangement of the cells marked 1, 2, 3 in fig. 10, 
and the character of the nucleus of "1" as seen in fig. n, at once 
suggest that u 1 " is the megaspore preparing for its first division, and 
that "2" and "3" are two sterile cells cut off in the heterotypic and 
homotypic divisions respectively. A comparison with figs. 12 and 13, 
which unquestionably represent two normal megaspores before the 
first division, indicates however that either this interpretation is not 
the true one or that the ovule was not normal. 

In the preliminary account (10) a stage was figured which seemed 
to indicate a very large number of megaspores. Although every prep- 
aration of the same age showed an almost identical structure, it has 


proved to be merely that characteristic of an ovule which is destined 
to be abortive. 

It is uncertain whether three or four potential megaspores are 
formed, but it is quite certain that only one becomes functional. 
As shown in fig. 12, the surrounding cells begin to disintegrate so 
quickly that the other cells derived from the functional mother cell 
cannot be recognized. There is some indication that it is not the 
lowest megaspore which develops, as is usually the case, but no defi- 
nite statement can be made. The wall of the mature megaspore 
is so thin that it can be distinguished only with great difficulty. 

In fig. 13 a cell immediately above the megaspore shows one of 


the "brown bodies" very distinctly, and a part of this cell is drawn 
on a larger scale in fig. 14. Fig. 13 indicates that polarity may be 
established in the embryo sac before the first division of the megaspore 
nucleus, and many other preparations show the same, but by no means 
all (compare figs. 12 and 15). 


4. The female gametophyte 5 

After the first division of the megaspore nucleus, the walls of the 
embryo sac and disintegrating mother cells can scarcely be distin- 
guished. Their thickness has been unavoidably exaggerated in 
fig. 15. In the preparation figured (the only one showing a binucleate 
sac), no polarity is evident, nor are the nuclei arranged quite parie- 
tally. After the next division, however, the four nuclei are imbedded 
in (or rather protrude inward from) a 1 
cytoplasm, and there is some indication of polarity, two nuclei being 
situated near the upper end and two near the lower end of the sac, the 
two pairs being symmetrically placed with regard to one another 
(fig. 16). This is distinctly interesting in view of the recently revived 
opinion that the embryo sac of angiosperms is derived from that of 
gymnosperms by a series of reductions, although it is to be noticed 
that certain "primitive" angiosperms do not show the polarity so 
characteristic of the vast majority. The growth of the embryo sac 
(prothallus) proceeds rapidly now toward the apex of the nucellus, 

n s As pointed out by Coulter and Chamberlain (2), the history of a gameto- 
phyte should start from the mother cell rather than from the spore, but practically 
it is more convenient to consider sporogenesis separately. 


the apical part becoming very narrow and closely resembling the tip 
of an advancing pollen tube. The breaking down of certain cells 
in the axis of the nucellus, facilitating the upward growth of the pro- 
thallus, has already been referred to. The divisions of the nuclei 
are doubtless simultaneous until at least 64 are present (exactly that 
number having been counted in serial sections), and possibly until 
wall formation begins. Fig. 17 shows in outline the general structure 
of the nucellus at the time when 64 nuclei line the embryo sac. Fig. 
18 is part of a tangential section of the prothallus after free nuclear 
divisions have ceased. The nuclei each contain a single nucleolus 
which can invariably be seen to be hollow. 

Cell formation takes places as in most other gymnosperms, and 
has already been described and figured (io,fig. 9). A large number 
of nuclear divisions have been seen in the "alveoli," certain features 
of which strongly recall the peculiar divisions described by Lawson 
(4) as occurring at this time in Cryptomeria. All the earlier phases 
(figs. 19 and 20) are quickly passed through, but that shown in fig. 21 
is very persistent, the marginal fibers having almost the appearance 
of a cell wall. The wall between the nuclei, however, is always 
developed, but in cases where its width does not nearly equal the diam- 
eter of the cell, it is just possible that wall and fibers may disappear 
again and thus give rise to a binucleate cell. It has not been demon- 
strated, however, that binucleate cells ever originate in this way, and 
the chief interest of fig. 23 is in providing a connecting link between 


cells of Cryptorti 
conifer prothalli. 

Very soon after cell divisions are complete, the archegonium 



than the surrounding cells and their nuclei are a little larger (fig. 22). 
The position of the archegonia ' is somewhat variable, but usually 
two distinct types seem to be formed: (1) a single group of about 



(2) one, or usually more than one, group of few or several archegonia 
(rarely a single archegonium) which are found near, but not abutting 
on, the upper part of the pollen tube, and deep-seated in the prothal- 

6 In the few cases where two pollen tubes reach maturity, a group of this kind 
is organized in connection with each tube. 


lus (fig. 23). It is certain that neck cells are formed in the former 
(1), although they are very evanescent and much more difficult to 
demonstrate than in the archegonia of Finns, for example; but they 
have not been demonstrated in the deep-seated archegonia (2), 
although it is not unlikely that this is merely due to the difficulty of 

from ot 
E Junip 

In a recently 

figured in which a small deep-seated archegonium occurred outside 
the normal group. The figure leads one to suppose that no neck 
cell could be demonstrated, but no statement on this point is made 
in the text. After fertilization no neck cells can be found in either 
type of archegonium, which accounts for the suggestion made in my 
former paper (10) that neck cells are never formed in Widdringtonia. 
The neck, where present, consists of a single tier of four cells (fig. 23). 
The central nucleus of the archegonium divides (in the first type 
of archegonium) simultaneously in the whole group (fig. 24), a con- 
spicuous spindle being organized with long, slender, and considerably 
twisted chromosomes (fig. 24) . The reduced number of chromosomes 
(six) may be counted fairly readily in this division. It is to be noticed 
in fig. 23 that although the central nucleus of every archegonium in 
the basal group is dividing, yet the nuclei of the other archegonia are 
still in the resting condition. The evidence that these do not divide 
is purely negative, but the possibility of the deep-seated archegonia 
being really archegonium initials only, must not be overlooked. 
The case shown in fig. 16 of my former paper (10) indicated that 
rarely a persistent ventral canal nucleus is formed, but out of scores 
of prothalli examined, containing thousands of archegonia, this is 
the only case in which such a state of things was clearly indicated. In 
no other preparation has the ventral canal nucleus been certainly 
identified, although the occurrence of the spindle which initiates its 
formation furnishes additional evidence that it must be formed. 
Doubtless it is normally very evanescent. From Strasburger's 
(12) figures of the same nucleus in Juniperus, it would seem that it 
is by no means a conspicuous object in that genus, and although 
Lawson (5) figures a large ventral canal nucleus in Thuja, it evidently 
disappears quickly, as it is not figured in the mature archegonium, 
and the same was found in Ottley's recent paper on Juniper us (9)- 




The formation of binucleate and multinucleate prothallus cells 
has already been described. It is initiated at about this time, but 
nuclear divisions continue to occur 
embryo formation. Two such divid- 
ing nuclei are drawn in 

during the earlier phases of 





The structure of the mature arche- 
gonium is much like that of other 
Cupressineae, with a large, centrally 
situated oosphere nucleus and a basal 
(or sometimes partly lateral) vacuole 
(fig. 28). The neck cells do not per- 
sist long, the archegonia of the basal 

group finally opening into the pollen 

The total number of archegonia 
of both kinds organized in a pro- 
thallus varies considerably. Some- 
times there are only about 25-30, 
usually about 40-70, and occasionally FlG . 1 — Microphotograph of a 

as many as IOO. The microphotO- tangential section of a prothallus in 

graph (text fig. 1) is taken from a which about I0 ° arche g° nia were 
tangential section of a prothallus in 



int. About 50 of these appear, 
cut transversely, in this section. A prothallus containing about 50 
in all was sketched in my preliminary paper (10, fig. 15). 

5. The male gametophyte 

The earliest stages of the germination of the microspore have been 
already described, but the description will be briefly repeated here. 
The mature pollen grain is uninucleate and there is no evidence that 
any prothallial cells are formed. Thus the "pollen grain" is here 
synonymous with the " microspore.' ' 
tion usually consists in the formation of a solid outgrowth from the 
exospore, w r hich applies itself closely to the surface of the nucellus, 
sometimes with a tendency to grow down between the nucellus and 
integument. About three pollen grains usually begin to germinate 

The earliest stage of germina- 


in each ovule, but generally only one and never more than two reach 
maturity. A little later the solid outgrowth of the exine becomes 
hollow, and the intine then grows through it, forming the tube, which 
at once penetrates a short way into the nucellus, the single nucleus 
passing into the tube. Occasionally a tube is formed at once on 

The time during which the pollen tube remains uninucleate seems 
to vary a good deal, a tube with a single nucleus having been found 
associated with a four-nucleate prothallus (fig. 29), while in two or 
three cases a tube containing three nuclei was found in an ovule 
before the first division of the megaspore nucleus; one such case is 
shown in figs. 30 and 31. No differentiation of cytoplasm can be 
seen here around the upper nucleus, doubtless the body cell nucleus, 
but the preparation is noteworthy as the only one in which a difference 
in size could clearly be seen between the two sterile nuclei, that nearest 
the apex of the tube (the "tube" nucleus) being considerably larger. 
Fig. 32 is a detailed drawing of the tube, the position of which is 
shown in fig. 17. Its size is approximately the same as that of figs. 
30 and 31, but the structure is somewhat different. The two sterile 
nuclei are precisely alike in every respect, as they are in every prep- 
aration in which they occur, except in the one noted above and in 
one abnormal tube referred to later. The body cell is here sharply 
demarkated from the surrounding cytoplasm in which the other nuclei 
are imbedded. 

It should be mentioned here that the occurrence of only two nuclei 
in the tube at a stage somewhat later than this, as previously reported 
(10), has not been confirmed in any other preparations. It is curious 
that, in fact, the binucleate stage has never been certainly identified, 
as it is not improbable that the case just mentioned is really a three- 
nucleate tube, and that defective preparation is responsible for the 
appearance of only two. It may be added that the preparation there 
figured is from material collected very soon after the work was begun, 
and before the best fixing agents and oven periods had been satis- 
factorily ascertained. On the other hand, it may be that the uninu- 
cleate condition had persisted later than usual in this case, and that 
only two nuclei are actually present. In any event, it is certain that 
the two sterile nuclei must be cut off in somewhat rapid succession. 


After they are formed, the pollen tube renews its activity and rapidly 
grows until it reaches the megaspore membrane. It is only rarely 
that the apex of the membrane is penetrated; more commonly the 
tube grows down a little farther before entering the prothallus. The 
entry is however almost invariably effected before the commencement 
of wall formation, the latter usually taking place only after the tube 
has taken up its final position. At the point where the pollen tube 
pierces the megaspore membrane, a slight constriction almost invaria- 
bly occurs, while the tip of the tube dilates a little to form a kind of 
vesicle just within the prothallus. Preparations are often met with 
in which the contents of the tube are just opposite the constriction, 
indicating a certain amount of difficulty in passing this point. Inside 
the prothallus the wall of the tube becomes thicker (fig. 33). When 
the difficulty of passing this point is overcome, the tip of the tube 
rapidly advances to about, or a little beyond, the middle of the pro- 
thallus. During the whole later growth of the tube the individuality 
of the body cell is less evident than at the stage shown in fig. 32, and 
its cytoplasm may be quite indistinguishable from the rest of the tube 
cytoplasm. A case of this kind, in which all three nuclei are imbedded 
in a common mass of cytoplasm, has already been figured (10, fig. 10), 
and is not uncommonly met with (see also fig. 33). 

Perhaps the most usual case is shown in figs. 34 and 35. Here the 
body cell is distinct, not only from the surrounding cytoplasm- of the 
tube, but also from the cytoplasm in which the sterile nuclei are 
imbedded. With the triple stain, the cytoplasm of the body cell 
stains red, that in which the sterile nuclei are situated takes the orange, 
w r hile the rest of the tube cytoplasm is a deep violet. In later stages 
the body cell increases very considerably in size. Its structure shortly 
before division has already been described and figured. Some 
uncertainty was expressed as to th e fate of the two sterile nuclei. It 
was suggested that they were probably absorbed by and became indis- 
tinguishable from the cytoplasm. Later preparations have in the 
main supported this view, but sometimes these nuclei are recogniz- 
able in connection with a quite mature body cell, in a prothallus in 
which wall formation is complete. 

Only a single preparation has been obtained showing the two male 
cells fully organized (figs. 36 and 37). Each cell is surrounded by 


a definite wall (or membrane) and is more or less hemispherical. The 
cytoplasm is dense and very homogeneous, and in the center of each 
cell is a large nucleus with no nucleoli. The entire absence of starch 
in the male cells is noteworthy, as compared with other Cupressineae. 
In this preparation the two cells are just beginning to separate, and 
probably they would finally become rounded off as reported by Law- 
son (4) in Libocedrus. In contact with one of these cells (see figs. 36 
and 37) is a curious body which presents the appearance of a vesicle 
containing a number of small, deeply stained granules. No light 
can be thrown on the morphology or functions of this body. No 
trace whatever can be seen of the two sterile cells. Possibly their 
disappearance may be connected w r ith the appearance of the vesicle, 
but this is merely conjecture. 

6. Fertilization and embryogeny 

The actual process of fertilization has not been seen in Widdring- 
tonia; probably it agrees essentially with that in other Cupressineae 
as has been found to be the case in Callitris. 

The first stage of the proembryo shows two free nuclei arranged 
lengthwise in the archegonium (fig. 38). The two nuclei are alike 
in size and structure. No starch is noticeable in the proembryo et 
this stage, but in all later stages seen, a considerable amount of starch 
is present. In later stages the structure has often been very difficult 
to interpret. Although a considerable number of preparations has 
been obtained, the sections have nearly always happened to be oblique. 

g method 

in order to get over this difficulty. Serial sections were drawn in ink 
on gelatin plates, the drawings being then placed together in sequence. 
If the thickness of the plates is to the thickness of the sections as the 
diameter of the drawing is to the diameter of the section, then a solid 
figure is obtained which represents accurately the object sectioned, 
the structure of which can be seen fairly clearly through the trans- 
parent gelatin. The gelatin plates can be easily stuck together by 
wetting them slightly. 

Figs. 39 and 40 are reconstructed from serial sections. They 
show proembryos containing five and ten nuclei respectively. * n 
fig. 39 delicate walls can already be seen between the nuclei, the 


appearance of which indicates that they probably arose as cleavage 
planes. In fig. 41 a transverse section of the upper four nuclei is 
shown. The presence of kinoplasmic radiations (spindle fibers ?) 
between these four nuclei indicates that probably they had a common 
origin and that after the first division of the oospore the basal nucleus 
remains undivided. It is evident that the early development of the 
proembryo does not show any resemblance whatever (after the first 
division) to that described by Lawson (5) for other Cupressineae 
{Thuja and Libocedrus) , and by Ottley (9) for Juniperus, where 
eight free nuclei are organized before walls are laid down. 

& iu xxv ^ iiwww ^ xv - vx & 

Fig. 40 shows diagrammatically the structure of a ten-nucleate 


It was not quite clear 

whether the walls extended to the upper three nuclei or not. The 


how the second form arose from the first. Probably there is some 
variation in the earlier divisions; compare, for instance, fig. 40 with 
the two proembryos figured in my preliminary account (10, fig. 16). 
Embryo development has already been briefly described. 

Fig. 42 is a drawing of a germinating seed before the cotyledons 
are withdrawn from the testa. Fig. 43 shows the upper part of the 
same seedling after removal of the testa. Often the testa is carried 
up on the tip of one of the cotyledons in germination. The seedling 
structure (anatomy) of Widdringtonia cupressoides differs somewhat 
from that recently described by Hill and de Fraine for other species 
(3), and an account of it has already been published from this labora- 
tory, together with a description of a remarkable twin seedling of 
the same species (Morris 8). 

In the plumular development a pair of opposite leaves succeeds 
the cotyledons, and is found in a plane perpendicular to that of the 
cotyledons. These leaves are followed by from about three to about 
ten alternating whorls of four leaves (text fig. 2). The structure and 
leaf arrangement of a tricotyledonous embryo have proved rather 
interesting. The three cotyledons are equal in size and have prob- 
ably been equivalent in development and are succeeded by alternating 
whorls of three primordial leaves. This is shown in the photograph 
(text fig. 3). The transition region is longer than in the normal 
seedling, and only at its lower end is there any indication that the 




three cotyledons are not all equivalent. Here two of the xylem groups 
grow somewhat smaller and approximate and finally join, thus giving 
the diarch root structure typical of the normal seedling. 

7. Abnormalities 

The tip of the pollen tube of fig. 23 contains a curious body cell 
which is shown more highly magnified in fig. 23a. Its appearance 

strongly suggests that it is 
not a normal body cell just 
after the division of the 
nucleus, but that during or 
immediately following the 
division disintegration of the 
nucleus has begun. 

In fig. 34 a slight irregu- 
larity is evident, inasmuch 
as a small fragment of the 

^:;v;;,.^.^;^; :; \::.^;;..v../>,^^.-;.;;: 

body cell nucleus has been 
constricted off and left just 

Fig. 2.— Diagrammatic cross-section of the behind in the Cytoplasm of 

cotyledons and plumule of Widdringtonia, show- the Cell. It is barely pOS- 

ing arrangement of the primordial leaves in a s j b , e ^ ^ have 

dicotyledonous seedling. 

been caused by defective 
preparation, but one other case has been seen in which a small papilla 
protruded in precisely the same direction from the body cell nucleus 
(not figured). Moreover, no such 
phenomenon has ever been seen by 
the writer in the case of any other 
large nuclei, either in Widdring- 
tonia or any other plant. 

One very curious abnormal male 
gametophyte is shown in fig. 44. 
Here the sterile nuclei are in the 
swollen tip of the tube (which is 

Surrounded by the USUal group of FlG - 3-— Photograph of a transverse 

archegonia), but the body cell has 
been left far behind. 

section of a tricotvledonous seedling- 


The nuclei are shown in more detail in figs- 
It is noticeable that the body cell does not exhibit such 


a distinctive structure as is normally the case, while one of the sterile 
nuclei is clearly larger than the other and is also distinguished struc- 
turally from it. This also is unusual and suggests the possibility that 
this larger cell might have eventually taken the place of the tardy 
body cell. 

The other two cases figured which seem to be abnormal are both 
taken from archegonia occurring in prothalli in which the pollen tube 
had lost its contents, but in which no proembryo was evident. Both 
might conceivably be taken to represent stages in the fusion of sexual 
nuclei. Fig. 45 shows an archegonium which was unhesitatingly 
considered to contain the sexual nuclei in contact, but on examination 
of these nuclei with a more powerful objective they were found to 
have the structure indicated in fig. 45a. The whole of the contents 
seems to consist of homogeneous and rather dense nuclear plasm 
except for the four nucleoli ( ?). Each of these nucleoli ( ?) consists of 
a membrane from the inside of which a plasmic strand has contracted 
on which are regularly arranged about six (three or four in optical 
section) very deeply staining bodies, of the shape and size figured. 
It is suggestive that the number of these bodies in each nucleolus ( ?) 
should be the same as the haploid number of chromosomes. It is 
also suggestive that two of these nucleoli (?), differing only in size, 
should be present in each of the fusing nuclei, if such they are. 

Figs. 46a and 466 are two successive sections of the same arche- 
gonium nucleus. The structure is so strikingly different in the two 
that it is difficult to believe they are actually parts of the same nucleus; 
but the fact is clear. In each section all the deeply staining structures 
(chromatin granules ? and nucleoli ?) have been figured, whether 
or not they occur in the same optical focus. 

Beyond the suggestion made above, I have no opinion to offer 
on these figures. Since they occur at such an interesting stage of the 
life history it seems desirable to place them on record. 

8. General 

Since much variation has been noticed among the Cupressineae 
in the time elapsing between pollination and fertilization, it is inter- 
esting to compare Widdringtonia in this respect. The chief point 
of interest is that although a very long period intervenes between 


pollination and fertilization, yet there is no break in the cQntinuity 
of development, such as probably occurs in those temperate conifers 
which are pollinated in one season and fertilized in the next. It is 
further interesting to notice that there are absolutely no fixed periods, 
so far as the writer can judge, when definite stages may be found. It 
may be not out of place to observe that these facts have made the 
working out of the life history very much more difficult than w r ould 
otherwise have been the case, which must be the excuse for the gaps 
which still remain in the present account. 



possibly early in February) of the present year are still far from ready 
for fertilization. They are still under observation and may probably 
be fertilized by the time this paper finally goes to press, in which case 
a postscript will be added. 

Very young female cones may be found at any time, but soon die 
off unless pollinated. The time of appearance of the male cones is 
very variable. During the present year the writer has failed to find 
a single male cone, except in the early part of January. Nevertheless, 
recently pollinated ovules were collected at the end of June, so that 
there probably must have been male cones in the vicinity during May. 

During 1908, male cones were collected in April and again in 
May (mature in both cases and separated by a full four weeks). 
A large number of trees were examined frequently and carefully 



time and in* one clump of trees may be of interest. About March 1 
the following female cones were found: (1) very young; scales still 
widely open; pollinated (probably about one month previously); 
(2) medium size; still quite green; ovules w r ith well-developed 
embryo sac; (3) full grown, brown, but with junction lines of scales 
still green; ovules with full-size embryo sac but no trace of cell for- 
mation; (4) full grown, uniformly light brown; integument discol- 
ored; young embryos; (5) dark brown, showing signs of dehiscence; 
mature seeds. It was afterward found that (3) could be segregated 
into two separate batches, in one of which fertilization took place 
about the end of April, in the other about the middle of September. 


As stated in the introduction, it is not proposed to discuss the facts 
here reported until the corresponding facts in the life history of 
Callitris are published, when the two genera may conveniently be 
compared and contrasted. 


The microsporophylls are arranged in decussate pairs and each 
bears four microsporangia. 

The mature pollen grain is uninucleate. 

The four equal megasporophylls are opposite and decussate. 

About 64 megaspore mother cells are organized at the base of the 
nucellus, but only one is functional Peculiar structures are noted 
at the poles of the non-functional megaspore mother cell. 

The megaspore may show polarity before the first division of the 
nucleus, or when four nuclei are present. The divisions of the embryo 
sac nuclei are probably simultaneous. 

Cell formation in the prothallus is normal in most respects, but 
certain peculiarities have been noted in the nuclear divisions which 
suggest comparison with Cryptomeria. 

The archegonia are never situated at the apex of the prothallus, 
but in several groups organized in relation to the pollen tube and 
deep-seated in the prothallus. The lowest group abuts on the pollen 
tube; the upper groups do not, and may possibly represent archego- 
nium initials. In the lowest group four neck cells are formed and a 
ventral canal nucleus is cut off. The total number of archegonia 
in a prothallus varies from about 30 to about 100. 

The microspore nucleus remains undivided for a long time. In 
other respects the development of the male gametophyte is more or 
less normal, but the so-called "stalk" and "tube" nuclei are almost 
invariably exactly alike and tend to disappear completely in the ma- 
ture pollen tube. 

The proembryo completely fills the archegonium, but the arrange- 
ment of the cells is somewhat variable. Walls are formed when less 
than eight free nuclei are present. 

The mature embryo has two (very rarely three) cotyledons. 

The cells of the mature prothallus are all binucleate or multinu- 


Stages in the life history are found not to correspond to definite 
seasons, and a long time elapses between pollination and fertilization. 

Botanical Laboratory 
South African College, Cape Town 

Postscript (March 26, 1910). — Wall formation in the prothallus 
of ovules pollinated about January, 1909, is now taking place. Fer- 
tilization is likely to follow in the course of a few days. It may there- 
fore be stated that an interval of fourteen or fifteen months will elapse, 
in this instance, between pollination and fertilization. 


1. Carothers, I. E., Development of ovule and female gametophyte in Ginkgo 
biloba. Bot. Gazette 43:116-130. pis. 5,6. 1907. 

2. Coulter, J. M., and Chamberlain, C. J., Morphology of spermatophytes. 
Part I. New York. 1901. 

3. Hill, T. G., and Fraine, E. de, The seedling structure of gymnosperms. 
I. Annals of Botany 22:689-712. pL 35. figs. 1-8. 1908; II. Annals of 
Botany 23:189-227. pi. 15. 1909. 

4. Lawson, A. A., The gametophytes, fertilization, and embryo of Cryp- 
tomeria japonica. Annals of Botany 18:417-444. pis. 27-30. 1904. 

5. , The gametophytes and embryo of the Cupressineae, with special 

reference to Libocedrus decurrens. Annals of Botany 21:281-^01. pis. 24- 


26. 1907. 


Normandie. Structure et affinities du 

Bennettites Morieri Sap. et Mar. Mem. Soc. Linn. Normandie 18: 
pis. 1-6. 1894. 
7. Masters, M. T., Notes on the genera of Taxaceae and Coniferae. J 

Linn. Soc. 30:1-42. 1893. 


and a brief account of the vascular system of the normal seedling. Phil. 
Trans. Roy. soc. S.A. 1 1411, 412. figs. 1, 2. 1909. 
9. Ottley, A. M., The development of the gametophytes and fertilization in 
Juniperus communis and Juni perns virginiana. Bot. Gazette 48:31-46. 
pis. 1-4. 1909. 

10. Saxton, W. T., Preliminary account of the ovule, gametophytes, and embryo 





former paper. It is not necessary to repeat the titles here, only those actually cited 
in the text being given. 


11. Saxtox, W. T., Contributions to the life history of Callitris. To appear in 
Annals of Botany 24: (July) 1910. 

12. Strasburger, E., Die Angiospermen und die Gvmnospermen. pi. 22. 



All figures were drawn with Zeiss camera lucida, microscope, and lenses, 
except figs. 39 and 40 (see text). In all: b, "brown bodies"; c, micropyle clos- 
ing cells; d, megaspore mother cells; e, functional megaspore mother cell; /, 
male cells; h, integument; n, nucellus; q, body cell nucleus; p, prothallus; 
r, resin cavity; s, sterile nuclei; t, pollen tube; v, vascular bundle. 


Fig. 1. — Transverse section of mature microsporophyll, showing positions of 
four microsporangia, resin cavity, and vascular bundle. X35. 

Fig. 2. — Transverse section through micropyle of an ovule about same age 
as fig. 3; the formation of the "wing" has only just begun. X205. 

Fig. 3. — Median longitudinal section of a young ovule, showing position of 
megaspore mother cells. X85. 

Fig. 4. — Section through megaspore mother cells in an ovule similar to fig. 3. 

Fig. 5. — Two non-functional megaspore mother cells, showing characteristic 
appearance of the "brown bodies." X825. 

Fig. 6. — Part of nucellus of an ovule slightly older than fig. 3, showing forma- 
tion of schizogenous cavities and sliding growth of three cells. X 205. 

Figs. 7, 9. — Megaspore mother cells in synapsis. X825. 

Fig. 8. — Position of mother cell of fig. 7 in relation to surrounding cells. X 205. 

Fig. 10. — Sporogenous cells at a somewhat later stage (see text). X500. 

Fig. 11.— Cell 1 of fig. 10. X825. 

Fig. 12. — A functional megaspore and some of the surrounding cells. X480. 
Fig. 13. — A functional megaspore and one of the surrounding cells. X480. 

Fig. 14. 


Fig. 15. — A binucleate embryo sac showing disorganization of other sporo- 
genous cells. X360. 

Fig. 16. — A four-nucleate embryo sac and the disorganizing mother cells 
surrounding it. X2o^. 

Fig. 17. 

position of poll 

and embryo sac when 64 nuclei are present in the latter; the region occupied by 
disorganizing spore mother cells is shaded. X35. 


Fig. 18. — Some prothallus nuclei in tangential section of the sac, showing the 
hollow nucleoli. X825. 

Figs. 19-21. — Three dividing nuclei in the "alveoli." X1240. 


Fig. 22. — Archegonium initial. X205. 

Fig. 23. — Part of a longitudinal section of a mature prothallus, showing the 
pollen tube, the basal group of archegonia, and three accessory archegonia; the 
nuclei of all the basal archegonia are dividing. X120. 

Fig. 23a. — The body cell of fig. 23. X825. 

Fig. 24. — Part of one of the archegonia of fig. 23, showing the spindle con- 
cerned in cutting off the ventral canal nucleus. X825. 

Fig. 25. — The neck of an archegonium in transverse section. X205. 

Figs. 26, 27. — Dividing nuclei in the prothallus cells. X150C. 

Fig. 28. — Mature archegonium; the position of the pollen tube is indicated 
above the neck. X360. 

Fig. 29. — Young pollen tube; only a single nucleus is present. X480. 

Fig. 30 — Tip of young pollen tube showing three nuclei. X825. 

Fig. 31. — Part of nucellus, showing position of the tube of fig. 30. X50. 

Fig. 32. — Young pollen tube with body cell and tw T o sterile nuclei; the posi- 
tion of this tube is shown in fig. 17. X825. 


Fig. 33. — Tip of pollen tube just entering the prothallus. X205. 

Fig. 34. — Tip of pollen tube after growth of the tube has ceased; the sterile 
nuclei with their surrounding cytoplasm form a distinct cell; a small fragment 
of the body cell nucleus has separated off. X205. 

Fig. 35. — The body cell nucleus of fig. 34. X825. 

Fig. 36. — -Tip of mature pollen tube, showing position of male cells and the 
relation of archegonia to the pollen tube. X205. 

Fig. 37. — Male cells of fig. 36. X480. 

Fig. 38 — Part of a transverse section of a prothallus, showing an archegonium 
containing two free nuclei, the daughter nuclei of the oospore. X360. 

Fig. 39. — Diagrammatic longitudinal section of proembryo with five nuclei; 
reconstructed from serial sections. X170. 

Fig. 40. — Similar section of proembryo w r ith ten nuclei; reconstructed from 
serial sections. X230. 

Fig. 41. — One of the sections from which fig. 39 was constructed, slightly 
oblique, showing the upper four nuclei and the cleavage planes (?) between 
them. X200. 

Fig. 42. — Young seedling. X 1 . 5. 

Fig. 43. — Cotyledons of same. X3.7. 

Fig. 44. — Abnormal pollen tube (see text). X50. 

Figs. 44a, b. — Nuclei of fig. 44. X 205. 

Fig. 45. — Abnormal archegonium. X205. 

Fig. 45a. — Part of fig. 45 (see text). X825. 

Figs. 46a, b. — Two adjacent sections of the nucleus of an abnormal arche- 
gonium. X825. 



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Lula Pace 

(with eleven figures) 

In September 1906 Miss S. M. Hague sent me some fern pro- 
thallia from a swamp in northern Indiana, where they were growing 
luxuriantly on rotten wood. Pieces of the wood were put into a 
glass jar, which was covered and placed on a table about seven feet 
from a window with a southeast exposure. They were kept very 
moist, yet watered with care, so as not to allow any water to get on 
the plants and thus cause fertilization. In spite of this precaution, 
however, enough moisture probably collected on the plants to permit 
fertilization occasionally, for sporophytes developed at intervals. 
Some of these, as well as many of the gametophytes, were used in 
class work. A few of the sporophytes are about 7 cm. high. They 
resemble Slosson's figures of Dryopteris spinulosa intermedia 
(Aspidium intermedium Muhl.). Certain peculiarities found in this 
material seem worth describing. Some of the gametophytes are 
still growing and will be watched for further developments. 


The prothallia were of the typical heart-shaped form, the larger 
ones being about 5 mm. long. Those on which sporophytes have not 
developed have continued to grow, and many spores, which had been 
lying for a long time in the rotton wood, have germinated. The old 
prothallia have grown to unusual size and taken queer shapes, many 
of them being 15 mm. long, a few 23 mm., and one even measuring 
37 mm. The most striking thing about them, however, is the peculiar 
forms developed. A hasty observation of one of these often gives 
the impression of a number of prothallia near together and over- 
lapping, for they often branch, apparently from any part of the plant, 
as in fig. 1, which is a diagram of one of the simpler cases. At one 
point near the margin of another plant 21 of these branches were 
counted, all 1-1.5 mm. long, and each bearing 6-10 antheridia. In 

49 ] [Botanical Gazette, vol. 50 






some cases filamentous processes grow out (tig. 2) ; this figure was 
made from fresh material, and the cell walls may not be accurate in 

Figs, i, 2. — Fig. 1, diagram of one of the simpler prothallia with several ganie- 
tophytic branches; fig. 2, sketch of a filamentous process with antheridia, from living 

every case. Occasionally a cylindrical process was seen, as described 
by Lang (2), but it might possibly have developed into the ordinary 
form, for some of the branches have slender bases. In several 





diagram (fig. 3), which represents a small prothallium 5 mm. long. 

margins, but shows no 

The outgrowth has 

antheridia along the 

archegonia; these were present, however, on the main prothallium. 


Figs. 3-6. — Fig. 3, prothallium 5 mm. long, with apical process bearing antheridia 
but no archegonia, which are found on the main prothallium; fig. 4, section through 
a very young gametophytic branch with apical cell; the branch is at the edge of the 
section and near an archegonium, the cross-section of the neck of which is shown; 
fig. 5, section through the basal part of a gametophytic branch, showing its relation 
to the main gametophyte; fig. 6, an antheridiai prothallium 2$ mm. long, with many 
archegonia but no antheridia except in the basal region. 

Sections of this prothallium show nothing unusual in structure or 
cell contents. 

A section of one of these branches at a very early stage shows its 


It comes from the edge 

of the archegonial cushion, the section showing also the neck of an 
adjacent archegonium. So far as the apical region is concerned, it 


looks like a normal gametophyte. The basal part of a somewhat 
older branch is shown in tig. 5, which is broader and does not seem 
so closely related to archegonia, although near the cushion. Appar- 
ently any cell or group of cells may be rejuvenated and initiate this 
branching. The cells in the neighborhood of fig. 4 are very large 
for cells in a position so near the apical region, and they contain 
unusually large food bodies, apparently oil globules; this may be 
a condition that leads to branching. 

Another type of gametophyte is shown in fig. 6. This prothallium 
does not branch and is 23 mm. long. There are few or no antheridia 
on prothallia of this type except near the basal region, but the arche- 
gonial cushion is unusually wide and thick, and has a broad row of 
archegonia along both margins, with more in the center. Trans- 
versely, these rows have 1-5 archegonia very closely crowded together. 
In one gametophyte 16 mm. long, there were found by actual count 
142 archegonia on one side of the cushion, with apparently as many 
on the other side; so that instead of the usual small number of arche- 
gonia, there were probably not far from 300 on this particular game- 
tophyte, and other gametophytes of this type. The production of 
archegonia apparently goes on indefinitely, for the prothallia still 

looked vigorous. 

Antheridia and archegonia 

A rather complete series of stages in the development of antheridia 
and archegonia was examined, but most of them resembled the usual 
type. Two late antheridial stages are shown in figs. 7 and 8, the 
former with the spermatogenous cells in mitosis, and the latter 
with sperms almost mature. There are apparently 16 chromosomes, 
though none of the figures are in condition to permit absolute accuracy 
in counting. As a rule, antheridia are found only on the basal region 
of old prothallia or on the branches, but occasionally one or two appear 
at any point on the gametophyte. In a few instances immature 
antheridia were found among mature archegonia. An unexpected 
feature in archegonial development is their appearance far back from 
the growing point, so that young archegonia are found among the 
old ones. A peculiarity several times noted was two archegonia 
with no wall between the egg cells, but with two complete necks; 
otherwise nothing unusual was seen in their development. 






A few prothallia were placed in water for a short time, and fer- 
tilization took place in the usual way, one or more sperms entering 
the neck and reaching the so-called receptive spot, where one sperm 
enters the egg. The sperm nucleus fuses slowly with the egg nucleus, 


Figs. 7, 8. — Fig. 7, antheridium with s per matoge nous cells in mitosis, showing 
approximately 16 chromosomes; fig. 8, antheridium with sperms almost mature. 

the fusion apparently being completed while both are in the resting 
condition. A few normal embryos in well-advanced stages were 
found. These must have been formed from the fertilized egg or 
have been developed parthenogenetically. 


A peculiar structure was found on the gametophyte which bore 
the antheridia shown in fig. 8, but it was found on the upper surface. 
The diagram (fig. 10) shows the relation of this body and the anther- 
idium to each other and to the archegonia. It is clear that there are 
only two possible interpretations of this structure: it must be an 





abnormal antheridium, or it is a sporangium; if it is the latter, this 
is a case of apogamy. The great difference in size is seen by com- 

Figs. g, 10. — Fig. 9, a sporangium-like structure from the upper surface of the 
gametophyte from which was drawn fig. 8; two layers of cells outside the fertile region, 
the inner layer suggesting a tapetum; each layer consists of many cells; fig- J°> < * ia ' 
gram showing the relation of figs. 8 and 6 to one another and to the archegonial region. 

paring the figures, which are drawn to the same scale. Among all 
the antheridia examined from this material, there is no such difference 
in size, the antheridium shown in fig. 8 being a rather large one. I* 1 

19 io] 



all the antheridia only one layer of wall cells is present, and these 
are few and rather definitely placed. But this structure shows two 


distinct layers (figs, g, io), except in one section, and this at only one 
place (fig. n). Here for the 
space of two cells the wall is 
only one cell thick, but these two 
cells might divide and make the 

two layers complete. The outer 
layer consists of many cells, which 
always contain chloroplasts, but 
the next layer contains chloro- 
plasts in a few cells only. The 
nuclei in this second layer are 
quite different from those in the 
interior, and in some of the cells 
there is a suggestion of the usual 
tapetal appearance. It does not 
have the long slender stalk char- 
acteristic of the normal sporangia 
of the higher Polypodiaceae, but 
it probably would have elon- 
gated somewhat before maturity, 
making it very similar to the 
stalks of the Osmundaceae. Con- 
sequently, this structure is quite 
unlike an antheridium, and is 
almost perfect in its sporangial 
characters. As there were no 

Fig. ii. — A small portion of the same 
sporangium-like structure shown in fig. 9, 
it COllld not be but from another section; this is the only 
determined whether the nuclei Place where one layer of cells appears out- 

contained the sporophytic or 


side the fertile region, and these two cells 

gametophytic number of chromo- complete, 
somes, but we should expect 

the sporophytic number to appear, especially since another pro- 
thallium bore a mature sporangium of the Osmunda, type, with a 
few spores still remaining in the sporangium. In the absence of 
mitotic figures, such a sporangium easily invites speculation. It 


may be an x structure, and the spores may have been formed in the 
usual way, so that the new gametophyte will have only \x chromo- 
somes; an interesting surmise in view of the fact that nearly related 


Or there 

may have been a doubling of the number and a subsequent reduc- 
tion at the formation of spores. The appearance of a sporangium 
upon a prothallium will be accepted as a case of apogamy. 


Lang (2), in his work on apogamy, grew prothallia from spores. 
Such prothallia, when kept dry, in direct sunlight, and watered only 
from below, developed leaves, roots, and ramenta on the prothallia 
themselves or on the cylindrical processes. The process continued 
as a leaf, or it produced sporangia, and tracheids were found both 
in the process and in other parts of the prothallium. When sporangia 
were found on a cylindrical process, tracheids were always present 
in the underlying tissue. Yamanouchi (7) reports very slow growth 
and few abortive archegonia in his material kept in bright light 

and dry air. 

My material was not in bright light and was given plenty of 
moisture, being kept in as nearly normal condition as possible, 
except for the entire absence of liquid water. Consequently, if 
this is apogamy, the lack of fertilization is apparently the only factor 
involved in its appearance here, for in this material archegonia were 
very numerous and were normal in every respect, and fertilization 
did take place when water was supplied. In this respect it seems to 
be like Marsilea, where Shaw (3) and Strasburger (5) found that 
if megaspores of Marsilea Driimmondii were isolated, and therefore 
fertilization prevented, parthenogenetic (apogamous) embryos were 

The gametophyte number of chromosomes is approximately 16. 
This number is very suggestive of Osmundaceae (Smith 4, Stras- 
burger 6, Yamanouchi 8), which is reported to have 16 chromo- 
somes in the spore mother cell. However, the young sporophyte 
does not resemble the mature form of Osmunda, and whether it 
resembles the sporeling I cannot tell, not being familiar with the 
sporelings of Osmunda. 


The sporophyte structures afforded no opportunity for determin- 
ing whether it contained the haploid or diploid number of chromo- 
somes. Yamanouchi found the haploid number of chromosomes in 
apogamous embryos, and concluded that the number of chromosomes 
is not the only factor which determines the characters of the sporophyte 
and gametophyte. Strasburger (5) found two kinds of megaspores 
in Marsilea Drummondii, some with the haploid and some with the 
diploid number of chromosomes. It would be expected that those 
with the diploid number of chromosomes would develop sporophytes 
without fertilization, as the gametophyte, and consequently the egg, 
has the diploid number already present. Farmer and Digby (i) 
found a vegetative fusion of nuclei in two forms, thus getting the 
sporophytic number of chromosomes without ordinary fertilization. 

The question of apogamy and the literature on the subject will 
not be discussed further at present, as it is hoped the material may 
furnish further evidence of this condition. 

It is a pleasure to express my obligations to Dr. Charles J. 
Chamberlain for advice and criticism during this work. 


Prothallia kept for three years in the laboratory in as nearly normal 
conditions as possible, except for the absence of liquid water, continue 
to grow, but develop peculiar forms and branching of various types. 

The sex organs continue to develop, antheridia being found occa- 
sionally on the main plant in all positions, but especially on the 
branches. Archegonia become very numerous, approximately 300 
having been found on one gametophyte. These not only develop 
in the apical region, but also far back among the old archegonia. 

Fertilization may take place whenever liquid water is present, 
as shown in several cases where gametophytes were placed in water 
and sectioned later. 

Apogamy is present in a sporangium-like structure which lacked 
the long stalk of the Polypodiaceae, but was not unlike the younger 
stages of the sporangium of the Osmundaceae. It had two layers of 


gion, the inner 

Baylor University 
Waco, Texas 





1. Farmer, J. B., and Digby, L., Studies in apospory and apogamy in ferns. 
Annals of Botany 21: 161-199. pis. 16-20. 1907. 

2. Lang, W. H., On apogamy and the development of sporangia upon fern 
prothallia. Phil. Trans. Roy. Soc. London B 190:189-238. pis. f-11. 1898. 

3. Shaw, W. R., Parthenogenesis in Marsilea. Bot. Gazette 24:114-117. 


4. Smith, R. \V., The achromatic spindle in the spore mother cells of Osmunda 
regalis. Bot. Gazette 30:361-377. pi. 1. 1900. 

5. Strasburger, E., Apogamy bei Marsilea. Flora 97: 123-191. pis. 3-8. 1907. 


6. , Ueber Reductionstheilung, Spindelbildung, Centrosomen, und Cilien- 

bildung im Pflanzenreich. Jena. 1900. 

7. Yamaxouchi, S., Apogamy in Nephrodium. Bot. Gazette 45:289-318. 

pis. 9, 10. 1908. 

, Chromosomes in Osmunda. Bot. Gazette 49:1-12. pi. 1. 1910. 


I. F. Lewis 

(with ONE figure) 

Periodicity in the production of the sexual cells of Dictyota dichot- 
omy has been described by Williams (8) for Bangor, Wales, and 

Plymouth, and by Hoyt (2) for Beaufort, North Carolina. At 
Bangor the sexual products are liberated at fortnightly intervals, the 
rudiments of sexual organs appearing a few tides before the least 
neap, and the mature gametes being liberated 3-5 tides after the 
greatest spring. In October, however, the time relations are reversed. 
At Plymouth the crops are later, as well as slower in maturing, than 
at Bangor, liberation occurring 7-12 tides after the greatest spring. 
At Beaufort only one crop a month is produced, the initiation of 
rudiments occurring the day before, or the day of, and liberation 
taking place six days after, the greatest spring tide. 

These striking differences in the behavior of what is pronounced 
to be the same species in different localities make it desirable that 
careful observations be recorded for this form in other regions than 
those mentioned, and especially in those regions where the tidal 
relations are different. Williams surmises that "the periodicity of 
the sexual cells is an hereditary character, and consequently may be 
expected to manifest itself in seas and habitats where there are no 
tides." Oltmans (5, pp. 487, 488) states: "Williams findet einen 
Zuzammenhang der Entwickelung und Befruchtung und mit den 
bekanntlich in Abstanden von 14 Tagen auftretenden 'Spring- und 
Nipptiden.' Da solche im Mittelmeer fehlen, werden erneute 
Untersuchungen hier die Dinge zu klaren haben." 

With these facts in mind, I took occasion, during March and 
April 1908, to make daily notes on the condition of the sexual plants 
of Dictyota at Naples. 1 The results of these observations are here 

1 I take this opportunity of expressing my thanks to the directors of the Smith- 
sonian Institution for the privilege of occupying a table at the Zoological Station in 
Naples, and to Dr. R. Dohrn and Dr. Lo Bianco for their cordial cooperation 
during my stay in Naples. 

59] [Botanical Gazette, vol. 50 


The range of the tides at Naples is very much less than at either 
Beaufort or Bangor, though it cannot be said that tides are wholly 
lacking at this point. The daily readings of the tide gauge near 
Naples, furnished by the Italian government, agree closely with 
the tides predicted by the U.S. Coast and Geodetic Survey. Extra- 
ordinary conditions of wind and weather, however, may affect the 
range of the tides very appreciably. For the period studied, the 
maximum daily range of the tides was 2.1 feet, the minimum 0.4, 
the average 1 . o. The water level varied from o . 7 feet below to 1 . 5 
feet above mean low water of spring tides. The following table 
gives an idea of the relative ranges of the tides at Bangor, Beaufort, 
and Naples. 



Bangor . . 
Naples . . 

Avenge range 

1 7 . o feet 

1 .0 

Difference in height 
of low water at 
spring and neap 

5 • 7 f eet 

The sexual cells of Dictyota were found to be produced at regular 
intervals, the time of initiation of the rudiments and liberation of 
the mature gametes bearing a definite relation to the periodic changes 
in the tides. The crops are borne, as at Bangor, at fortnightly 
intervals. Initiation of the rudiments occurs on the same day as 
general liberation of the mature gametes, this being two or three 
days after the least neap tide. The number of days required for 
the development of a single crop is approximately sixteen. The 
development of the sori is fairly uniform, not being accelerated at 
the time of the spring tides. 

A comparison of the facts of periodicity in Dictyota at Bangor 
Beaufort, and Naples is given in the following tables. 

The accompanying chart (fig. 1) gives in graphical form the 
record of one crop from initiation to liberation, showing the relations 
between the tidal changes and the fructification of Dictyota. The 
chart should be compared with the charts given by Hoyt (2,pP- 
386, 387) in order to compare the behavior of Dictyota at various 





At any given time all the individual plants, collected at various 
points from Capo Miseno to Santa Lucia, are at approximately the 
same stage of development. There are noticeable differences, how- 
ever, as for instance in the plants collected on April 10. According 
to the chart, this date shows the beginning of liberation of gametes, 






A few days before the the 

least neap 
Day before or day of 

greatest spring 
2-3 days after least neap 


3-5 tides after greatest 

6 days after greatest 

2-3 days after least neap 

Time of develop 

12-17 days 
8-9 days 
16 days 


History of a single crop 









I 3 


Undivided rudiments 
8-16 cells in surface 
view of antheridia 
32-64 cells 
64 cells 

General liberation 
Belated sori 


Undivided rudiments 
Undivided rudiments 

Undivided rudiments 
Undivided rudiments 


2-4 cells 

Many sori. 64 cells 

General liberation 
Belated sori 

Undivided rudiments 
Undivided rudiments 

or 2-4 cells 
2-4 cells 
2-4 cells 
2-4 cells 
4 cells 
4-8 cells 
8-16 cells 
16 cells 
16-32 cells 
32-64 cells 
64 ceils 

General liberation 
Belated sori 

and also the beginning of initiation of rudiments. As a matter of 
fact, the numerous individuals examined on this day showed con- 
siderable differences among themselves. The male plants may be 
grouped in the following categories: (1) no antheridia empty, new 


crop not visible; (2) none empty, new crop inconspicuous; (3) 
few antheridia empty, new crop conspicuous; (4) antheridia empty 
near base of plant, nor. empty near apex, new crop barely visible; (5) 


4-celled in surface view; (6) antheridia nearly all empty, new crop 




barely visible; {7) antheridia all empty, new crop showing 4 cells. The 
majority of plants collected on this day were in the fourth or fifth stage. 
The tendency toward periodicity in the production of sexual 
cells is probably a hereditary character. That periodicity is not a 
series of simple responses to successive stimuli is shown by the obser- 
vation of Williams that individuals removed from the influence 
of the tides continue to show the usual periodicity, and by Hoyt's 
statement that individuals produced by vegetative multiplication 

2829&3I 4 \2 3 

5RRJ NG ! ! ' 



\ I iNEAP l i s Spring: I 1 NEAP 


1 ', ; fwsf i ; i S full j \ \ . ; last J i 

j ; ; QUARTER ! { j j . ! MOON j J • J ; QUARTER , | j 

\\'\ -H ■■{ | I ! ! ! i i i i ! ! I [ i I f i 

i ill. ' ill * ■ iii 




Fig. i. — Chart showing tidal relations at Naples from March 28 to April 27, 1908, 
with record of crops of Dictyota for the same period. The tide records were furnished 
by the Italian government, through the kindness of Dr. R. Dohrn, of the Zoological 
Station at Naples. The chart is to be compared with similar charts for Bangor, 
Wales, and Beaufort, N.C. (Hoyt 2). The zigzag lines show the rise and fall of the 
tide and the height of the water in relation to the mean low-water mark of spring 
tides. The curved lines show the development of the sexual crops (male) of Dictyota 
for the respective periods with their relations to the tides. The numbers opposite 
the crop curves indicate the number of cells seen in the surface view of each antheridium. 

and never subjected to the influence of the tides show approximately 
the same periodicity as those in their natural habitat. The former 
observation was confirmed by me at Naples, and goes to show that 
the habit of periodical reproduction is fairly well fixed. The habit, 
however, must have had its origin in response to external conditions, 
and it is a matter of considerable interest to ascertain, if possible, 
what factor or factors gave rise to the periodicity. The only sugges- 
tion made thus far is that of Williams that the effective factor is 
the increased illumination during low water of spring tides, a view 
from which Hoyt dissents. 

Williams' hypothesis seems inadequate to explain the periodicity 
at Naples. Dictyota flourishes there at a depth of many feet below 


the surface, and the difference between the height of low water at 
spring and neap tides is only 0.25 feet. This slight difference can 
hardly cause any considerable variation in the total illumination of 
the plants under consideration, not as much as is caused by alter- 
nating cloudy and sunny days. It is interesting to note, however, 
that both initiation and liberation occur at Naples on the day that 
low water occurs at or nearest midday. Thus low water at midday 
occurred at Naples on March 29, April 10, and April 27, 1908, the 
days when initiation and liberation were found to occur. Whether 
this is more than a coincidence is still to be seen, but the fact remains 
that the critical points in the sexual life of Dictyota coincide exactly 
with the periods of maximum intensity of illumination. 

If one considers the behavior of Dictyota at Naples alone, it seems 
a fairly satisfactory hypothesis that the effective factor in producing 
periodicity is the stimulus of the maximum intensity of light. When 
one comes to apply this hypothesis to the other regions where Dictyota 
has been studied from this standpoint, it becomes evident that, if 
true at all, the hypothesis is modified by other factors. At Beaufort, 
for instance, low water at midday occurs two days before new and 
full moon, while initiation occurs on the day before or the day of 
the greatest spring tide. At Bangor, on the other hand, low water 
at midday occurs about five days before new or full moon, and initia- 
tion takes place one to two days before the least neaps. At Plymouth 
low water of the greatest spring tides occurs at midday, and here the 
times of initiation and liberation coincide more nearly with the periods 
of neap tides. It is obvious, then, that the simple explanation that 
might suffice for Naples is not sufficient for other localities, and that 
the operative factor or factors must be sought by further investi- 

It seems possible that Dictyota, in adapting itself to differing 
conditions at various localities, has acquired its habit of periodicity 
in response to different factors. That a similar stage in the repro- 

duction of other algae may be induced by different stimuli has been 
shown by Freund (i) to be the case in Oedogonium and Haema- 
tococcus. In these forms the external conditions leading to the 
formation of zoospores differ according to the condition of the indi- 
vidual plants. For instance, cysts of Haematococcus form zoospores 


(i) when transferred from foul to distilled water, (2) when brought 
from darkness into light, (3) when water is replaced by sugar solution. 
Klebs's investigations (3) illustrate the same point for other algae 
and fungi. Since the fructification of other forms may be induced 
by different factors, it is possible that the same may be true for Die- 
tyota in various situations, and that the factors concerned in inducing 
periodicity may vary with the locality. 

Other organisms, both animals and plants, show a periodicity 
in the release of sexual cells comparable to that of Dictyota. In 
Amphitrite ornata, an annelid worm, "practically all sexual products 
are deposited within three days of spring tides'' (Scott 6, p. 332). 
Kuckuck finds (4) in Nemoderma tingitana that the gametes are 
released at times of neap tides. Tahara shows (7) that the oospheres 
of Sargassum enerve are released at fortnightly intervals at about the 
time of new and full moon. Other species of Sargassum are said 
by Tahara to show a similar periodicity, in which liberation occurs* 
at various intervals after the greatest spring tides. 

It is obvious, therefore, that periodicity in the release of sexual 
cells is a widespread phenomenon, probably to be attributed to 
various factors in different species, and perhaps to more than one 
factor in the same species in different localities. 

Randolph-Macon College 

Ashland, Va. 



Freund, H., Neue Untersuchungen iiber die Wirkungen der Aussenwelt auf 
die ungeschlechtliche Fortpflanzung der Algen. Flora 98:41-100. 1907. 

2. Hoyt, W. D., Periodicity in the production of the sexual cells of Dictyota 
dichotoma. Box. Gazette 43:383-392. 1907 

3. Klebs, G., Die Bedingungen der Fortpflanzung bei einigen Algen und 
Pilzen. Jena. 1896. 

4. Kuckuck, P., Neue Untersuchungen iiber Nemoderma. Wiss. Meeresunters. 
Abt. Helgoland N.F. 5 : 1904. 

5. Oltmanns, F., Morphologie und Biologie der Algen 2:59. Jena. 1905. 

6. Scott, J. W., Some egg-laying habits of Amphitrite ornata Verrill. Biol. 
Bull. 17:332. 1909. 

7. Tahara, M., On the periodical liberation of the oospheres in Sargassum 
(prelim.). Bot. Mag. Tokyo 23:151-153. 1909. 

8. Williams, J. Ll., Studies in the Dictyotaceae. III. The periodicity of the 
sexual cells in Dictyota dichotoma. Annals of Botany 19:531-560. 1905. 



Fungous diseases of plants 

Owing to the existence of an extensive system of experiment stations in the 
United States, one of whose chief activities has been the investigation of plant 
diseases, the conditions for the accumulation of facts relating to plant pathology 
have been unusually favorable. Following the progress made in investigation, 
the teaching of plant pathology has begun to develop chiefly in the agricultural 
colleges associated with the stations. Thus far, however, there has been no attempt 
to organize into a comprehensive text the vast material accumulated by plant 
pathologists and to make it available for teachers, although a need for such work 
has been felt by those who have attempted to teach the subject. The appear- 
ance of Duggar's book 1 on plant diseases is therefore both timely and desirable. 
The work, as the preface indicates, is designed primarily as a textbook, but its 
possible service as a reference book has also been kept in view. Aside from the 
brief historical introduction, it falls into three parts: (i) culture methods and 
technic, (2) physiological relations, and (3) fungous diseases of plants. 

The first part is designed to introduce the student to the methods and manipula- 
tions used in the study of fungous diseases. In it are treated the methods of han- 
dling apparatus, the preparation of culture media, the cultivation of organisms, 
and microscopical technic. On the whole, the directions are clear and to the 
point, and embody many details of manipulation which are acquired only through 
intimate experience with such work. The use of the freezing method for cut- 
ting sections should perhaps have been mentioned, especially as its adaptability 
for certain kinds of work has been recently emphasized. 2 

The part on physiological relations comprises a discussion of the germination 
of spores and the modes of life and relation to environmental factors of parasitic 
fungi, together with chapters on artificial infection, disease control, and the 
preparation of fungicides. It stands for the whole field which belongs peculiarly 
to the general subject of plant pathology. Considered from this standpoint, the 
treatment is surprisingly brief and much that is in the chapter does not belong 
there. This is especially true of that part of the chapter on germination which 
deals with methods, part of the section on environmental factors, and nearly all 
of the chapter on artificial infection. These should have been included in the 

1 Duggar, B. M., Fungous diseases of plants. 8vo. pp. xii + 508. figs. 240. Bos- 
tun: Ginn & Co. (undated). 

2 FREEMAN, E. M., The ether freezing microtome in botanical work. Science 
X.S. 25:747-749. 1907. 




first part on methods and technic. The field represented by this part has been 
so greatly enriched in recent times, by the addition of both facts and ideas, that 
the treatment seems wholly inadequate. 

The last part, making up the bulk of the work, deals individually with the 
plant diseases induced by fungi, under which the author includes the myxomycetes 
and bacteria. The arrangement is in the order of taxonomic sequence, each 
chapter representing one of the large divisions of the fungi, as Phycomycetes, 
Ascomycetes, etc. The individual diseases are treated in numbered sections of 
the chapters. A section is given to the discussion of each disease, except those of 
minor importance, which are grouped together. The arrangement serves to bring 
out the morphological relations of the disease-producing fungi, without laying too 
much stress on purely morphological and taxonomic features. Some clearness 
would have been gained if the discussion of orders and families had not been 
forced into the system under headings coordinate with those under which the 
individual diseases are discussed. The treatment of the diseases is clear and 
comprehensive, each being discussed with reference to its distribution, the influ- 
ence of environmental factors on its occurrence and prevalence, the life history 
of its causal organism, and the methods of its control. In the relative prominence 
given to the various diseases, the author has been guided by their economic impor- 
tance, but the scope has been made broad enough to include all of the common 
diseases injurious to cultivated crops. A few diseases not occurring in this country 
have been included, apparently for the sake of completeness. As a rule, the text 
is conservative and free from innovations at variance with current usage. There 
is one notable exception, however, in the introduction of a series of newly com- 
pounded terms to apply to certain artificial groups of the rusts, based on the num- 
ber of spore-forms present in the cycle. Thus we have "euautouredo" to include 
all autoecious rusts possessing all spore-forms, and "opsiautouredo" to include 
all autoecious rusts lacking the uredo stage, etc. Aside from any criticism that 
may be offered on account of the faulty composition of these unwieldy terms, 
the pedagogical soundness of introducing them for the first time through the 
medium of a textbook may well be questioned. 

In matters of detail, the work shows an unusual lack of care in the preparation 
of the manuscript or in proofreading. The following examples serve as illus- 
trations: on page 54 "Lowitz" is printed for "Lowit"; on page 86 "Scot" for 
"Scott"; and on page 121, second citation, "Histology" for "History." In 
the legend of fig. 50 a germinating oospore is described as a "germinating oogo- 
nium." On pages 272 and 276 "Von Schrenk" should read "Von Schrenk 
and Spaulding." On page 337 the citation of McAlpine should read "Stone- 
fruit trees" instead of "Stone fruits." Phoma Betae is said not to occur in the 
United States (p. 344), but it has been reported from Colorado and Kansas. 5 
On page 386, note, Arthur is said to have introduced the terms "pycnium, aeciuM, 

3 Orton, W. A., Plant diseases in 1907. Yearbook Dept. Agr. K)oj:s77'S^9't 
see also Yearbook 1906:502 {Phyllosiicta Betae Oud.). 


uredinium, and telium in substitution for teleuto, uredo, aecidial, and sperma- 
gonial stages" of the rusts, instead of the reverse order. On page 467 Trametes 
Pint is said to be the " chief cause of loss among fungi." 

On the whole, the book is an excellent presentation of the subject of plant 
pathology from an American standpoint. Most of its shortcomings relate to 
individual or minor details. In it the vast amount of material collected through 
the agencies of the experiment stations and the U.S. Department of Agriculture 
has been brought together for the first time in an easily available form. The 
facts presented are largely derived from American work and apply to American 
conditions. It is sufficiently comprehensive for a textbook, and will be of much 
service as a reference book in the field which it represents. The style is clear and 
concise, and the arrangement is that which the teacher would naturally adopt. 
The free citation of literature is of great service to both student and teacher. 
The book is abundantly illustrated, and both illustrations and press work are all 
that could be desired.— H. Hasselbring. 

The morphology of plants 

The third and last volume of Velenovsky's textbook* on the comparative 
morphology of plants deals with the flower of phanerogams, the ovule, pollination, 
embryo, seed, fruit, and the evolution of plants. Fertilization, parthenogenesis, 
and polyembryony are treated under the section on the ovule, preceding the 
description of pollination. The volume opens with the following definition of a 
flower: "The flower of phanerogams is a shortened axis of limited growth, which 


We are assured 


that this definition applies to all cases except the female structures of the genus 
Cycas, which are not regarded as flowers. 

The book deals almost entirely with the grosser external features of plants, 
e attention being given to the details of development. It must be confessed 
that the phase of morphology represented by this book is somewhat neglected by 
modern morphologists, who are likely to pay insufficient attention to the taxonomic 
side of botany. Morphologists should find the work useful as a reference and as a 
supplement to their taxonomy; but as a complete textbook of morphology it is 
not comprehensive enough to meet modern demands.— Charles J. Chamberlain. 



The cretaceous plants of Japan, s— This interesting product of the Anglo- 


4 Velenovsky, Jos., Verglekhende Morphologie der Pflanzen. Vol. III. pp. 478- 
pis. 6-g. figs. 400. Prag: Fr. Rivnac. 1910. For review of vols. I and II see Bot. 

Gazette 44:310. 1907. 

s Stopes, Marie C, and Fujii, K., Studies on the structure and affinities of 
cretaceous plants. Phil. Trans. Roy. Soc. London B 201:1-90. pis. i-Q. 1910. 


Japan. Only in rare instances were the authors able to make out the external 
form of the material studied, and in no case do they seem to have been able to 
correlate it with th© extremely abundant cretaceous genera known from impres- 
sions. To this initial disadvantage is added a not entirely satisfactory familiarity 
with the anatomical structure of living angiosperms and conifers. The eighteen 
species described as new in the memoir are consequently in some cases not really 
new, since they represent the parts of plants already known from impressions and 
recently identified structurally by American paleobotanists. In other instances 
the anatomical characterization is too vague and indefinite for subsequent use. 
In spite of these drawbacks, the memoir under discussion must rank as one of 
the most important recent contributions on the cretaceous flora, and it is much 
to be desired that the authors may be able to continue their investigations as they 
promise to do. 

Of the eighteen species described, four are cryptogamic, one being a fungus 
and three others ferns. An interesting cycad-like leaf, Niponophyllum, is described 
which differs from the leaf structure of living cycads in the complete absence of 
centrifugal wood, all the xylem being of the cryptogamic centripetal type. 

Of the other gymnosperms described, the most interesting is Yezonia, which 
is considered by the authors to represent a new genus, and of which they state 
"it is impossible to find any family among the gymnosperms with which we can 
satisfactorily include this plant." This view of the matter will hardly stand, 
since in every detail of structure it corresponds absolutely with Brachyphyllutn, 
the commonest conifer of the later Mesozoic, which, moreover, on anatomical 
grounds has been recognized recently as an araucarian conifer. Another gymno- 
spermous branch is also described under the new r generic name Cryptomeriopsis. 
Of this it may be stated that the description given of its internal organization by 
the Anglo-Japanese authors parallels with fidelity, so far as it goes, that of Geinit- 
zia Reichenbachi, recently described structurally from the North American Cre- 
taceous. Two imperfect coniferous cones are likewise characterized, Yezoslrobus 
and Cunninghamioslrobus. One Araucarioxylon and two species of Cedroxyloit 
complete the list of coniferous remains. 

Either as the result of a bad condition of preservation, or a failure to realize 
clearly the importance of detailed description, four angiospermous ligneous 
genera, all considered to be new (Jugloxylon, Populocaulis, Fagoxylon, Sabio- 
caulis), are insufficiently characterized. The detailed structure of the rays, the 
characters of the vessels and wood fibers, as well as the distribution of wood 
parenchyma, all important features in the description of angiospermous fossil 
woods, are entirely or almost entirely omitted. If the omission is due to faulty 
preservation, the woods are scarcely worth publishing. The genus Saururopsis 
is somewhat more clearly characterized. One genus (Cretovarium) representing 
a tricarpellary ovary is likewise described, but as the accompanying vegetative 
organs and even any considerable part of the floral apparatus itself are absent, 
it seems impossible to arrive at any satisfactory conclusion as to its affinities. 
One curious and unfortunate omission throughout the memoir is the almost com- 



plete failure to indicate the magnification used in the figures. This makes com- 
parisons on the part of other workers difficult or even impossible. 

In spite of the exceptions taken in various respects to the work of the Anglo- 
Japanese authors, it must be conceded that their line of investigation is one of 
great promise, and it is to be hoped that they will feel encouraged to continue it 
with a greater attention to definiteness in anatomical characterization.— E. C. 

Vascular anatomy of Gleichenia. — Boodle and Hiley, 6 from the study of 
the anatomy of Gleichenia pectinata and allied species, reach certain theoretical 
conclusions as to the origin of the tubular medullated stele. They report the 
result of the examination of the node and internode, as well as the branching stem, 
of certain species of Gleichenia, particularly G. pectinata. It is not surprising that 
they reach substantially the conclusions which have been published already by the 
senior author in earlier contributions. The published results in this case, how- 
ever, appear to indicate a certain modification of the position originally held by 
Boodle, to the effect that in all cases the pith is a part of the stele and is 
not derived by inclusion of the fundamental or ground tissue from outside the 
central cylinder; for the authors in this article use the term solenostelic, borrowed 
from Gwynme-Vaughan and employed by him in the sense of a tubular stele 
with internal as well as external phloem and inclosing fundamental tissue as a 
pith (a meaning stated by Gwynnt;-Vaughan himself to be equivalent to the 
reviewer's siphon ostelic with internal phloem). Although the English writers 
in this instance concede apparently the arrival of the solenostelic condition as the 
final result of the modification of the pithless protostele, they express the opinion 
that the pith appears first as the result of the transformation of some of the tracheids 
into a central mass of parenchyma, a condition followed by the appearance of 
ramular gaps in the stele as the result of branching, leading to the intrusion of 
phloem from the outside of the stele and ultimately of the fundamental tissue 


itself. Only at the end of the process do the leaf gaps appear and become patent. 
These views are all the more remarkable because in the same article the 
authors concede that the islands of parenchyma occurring in the petiolar strands 
of certain representatives of the Gleicheniaceae are derived from the cortex by 
inclusion, and were originally surrounded both by internal phloem and internal 
endodermis. The condition in which the included parenchyma is separated 
from the vascular tissues of the petiole by neither endodermis nor internal phloem 
is a result of progressive degeneracy. It appears almost an extreme example of 
the perversity of the human mind to explain the occurrence of central parenchyma 
in the leaf trace in a node diametrically opposite to that adopted for the appear- 
ance of a pith in the vascular tissues of the stem. If the fundamental tissues may 
be included in the leaf trace, there appears to be no reason why a similar process 
should not lead to the formation of pith in the axis. The adoption of this hypoth- 


Annals of Botany 23:419-432. pi. 2Q. 1909 



young individuals of various examples of vascular plants, a condition which finds 
no elucidation whatever in connection with the views adopted almost universally 
by English anatomists. — E. C. Jeffrey. 

Jurassic woods. — Gothan? describes a number of fossil woods from the 
Jurassic of King Karl's Land, none of which he refers to the Araucarineae. 
Two species, Phyllocladoxylon sp. and Xenoxylon phyllocladoides, are considered 
to belong in the region of the Podocarpineae. One Cupressinoxylon is also 
figured, which is compared with the C. McGeei of Knowlton. Next are two 
Cedroxyla, C. cedroides and C. transiens. To the latter of these he attaches much 
importance because of the presence of araucarian pitting together with the ray 
structure of the Abietineae. The author assumes that he has to do in this 
instance with an abietineous wood, in which certain araucarineous characters 
indicate the derivation of the Abietineae from an araucarian stock, in accordance 
with the conventional and generally received views. It does not seem to have 
occurred to the author that he may have an araucarian conifer with indica- 
tions of transition toward the Abietineae. The genus Araucariopilys, recently 
described by the reviewer, has the same ligneous characters as Gothan's Cedro- 
xylon transiens, only with a more pronounced resemblance to the Abietineae. 
It is nevertheless unquestionably an araucarian conifer. One abietineous wood 
is described, namely Protopiceoxylon extinctum. The wood in question has 
normally only horizontal resin canals, but vertical ones may occur as the result 
of injury. The author regards his wood as evidence that the vertical type of 
resin canal is older than the horizontal, forgetful that Goeppert and Penhallow 
have described much older Pityoxyla from the Carboniferous and Permian 
respectively, which show only horizontal canals. It seems, accordingly, that "Pro- 


topiceoxylon" with a much greater degree of probability, represents a stage in 
degeneracy toward the Cedroxylon type, rather than a primitive abietineous 
type antedating Pitfoxylon. The author lightly sets aside the evidence adduced 
on comparative anatomical and experimental grounds by the reviewer for the great 
age of the pinelike Abietineae. With admirable Teutonic frankness, he char- 
acterizes the reduction hypothesis of the phylogeny of the Coniferales as one to 
which, "wohl niemand beipflichten kann." He apparently loses sight of the 
fact that the reduction hypothesis is now quite generally adopted by competent 
morphologists for such degenerate groups as the Equisetales, Lycopodiales, etc. 
Had the literature on the fossil conifers, which has recently been published by 
American paleobotanical writers, been available, in all probability Gothan s 
opinion in regard to the fallacy of the reduction hypothesis in connection with the 
Dhylogenetic development of the conifers would have been expressed with con- 
siderably more reserve. — E. C. Jeffrey. 

7 Gothan, W., Die Fossilen Hoelzer Koenig Karls Land. Handl. Kgl. Svensk. 
Vetensk.-Akad. 42: no. 10. 


Chemotropism of roots. — In a preliminary paper, Porodko 8 reports upon 
the chemotropism of the roots of Lupinus albus and Helianthus annuus. Roots 
35 mm. long were placed in a lamella of agar varying in thickness from 6 to 60 mm., 
which separated the solution used from water. In all, 50 chemical substances 
were used, the concentration of which varied from o.m to o.ooiw. As a rule, 
the roots did not remain straight, but bent against or with the diffusion stream. 
The range of concentration between maximum and minimum depended upon the 
substance used and the thickness of the agar lamella. Concentrations close to 
the maximum caused bending against the diffusion stream or positive response, 
which effect was observed with both electrolytes and non-electrolytes. Porodko 
considers this a traumatropic response, due to the inhibition of growth on the 
up-stream side of the root. With lower concentrations, electrolytes and non- 
electrolytes affect the roots differently. The former cause great regularity as 
regards the direction of bending of the root, while the latter produce positive, 
negative, and intermediate responses. Acids, alkalies, and carbonates cause 
positive, and neutral salts negative bendings. The responses due to H and OH 
ions are considered to be traumatropic. The amount of negative response seems 
to depend upon the cation, being greater in the presence of one with a double 
charge than in the presence of one with a single charge. In many cases the 
responses are not all of one kind. Nevertheless, it is necessary to explain the 
cause of all. From his experiments, Porodko concludes that positive but not 
negative responses can take place in decapitated roots, and that the latter, but 
not the former, show up as after-effects, although only on the clinostat. The two 
reactions are different in nature, the positive being passive and caused by the 
inhibitory effect of the greater concentration on the growing region on the up-stream 
side of the root, the negative being active and due to the chemotropic effect of the 
diffusion stream, which tends to accelerate the growth on the up-stream side. 
Hence, upon the growing region of a root of Lupinus albus subjected to the influence 
of the diffusion stream of a chemical substance, two antagonistic tendencies are 
at work, the direction of bending of the root being dependent upon the relative 
strengths of the two tendencies. Roots of Helianthus annuus act differently 
from those of Lupinus albus, in that they show only traumatropic response, but 
why this is true is not known.— R. Catlix Rose. 

National Academy of Sciences. — At the annual session of 19 10 two botanical 
papers were presented (April 19), which may be outlined as follows: 

"The distribution of Agave in the West Indies," by William Trelease .— 
Three main types of Agave are recognized in the West Indies: one confined to the 
southwestern Cuban region, another to the Inaguas, and the third ranging through 
the entire archipelago. Subtypes of the latter are limited respectively to the 
Greater Antilles, the Bahamas, the Caribbees and the Leeward Islands, and the 

8 Porodko, Theodor, Ueber den Chemotropismus der Wurzel. Ber. Deutsch. 

Bot. Gesell. 28:50-57. 1910. 


adjoining Venezuelan coast. Within these groups specific differentiation is 
observable, so that each island isolated by a ioo-fathom channel has its endemic 
species, the islands with a common coastal plain possessing little if at all differ- 
entiated forms. The almost entire absence of the genus from South America 
and the geographic grouping of species and superspecies in the West Indies indi- 
cate that Agave penetrated from the Central American mainland, where it centers, 
and overran the terrain before the disruption into islands, two or perhaps three 
parent stocks being involved. 



Among the various vascular structures of gymnosperms that have been used to 
suggest progressive evolutionary changes, the vascular plate of the cotyledonary 
node is perhaps as significant as any, especially its connections with the cotyledons. 
Series of cycads, of conifers, and of other gymnosperms were shown to illustrate 
the following general tendencies: to reduce the cotyledons to two, to reduce the 
protoxylem poles of the vascular plate (and hence the root poles) to two, to elimi- 
nate certain vascular connections of the cotyledons, and to restrict the branching 
of strands within the cotyledons. 

Differentiation among chromosomes. — Crepis virens seems to afford promis- 
ing material for the solution of several difficult cytological problems. Juel had 
already found the diploid and haploid numbers in C. tectorum to be 8 and 4. 
Rosenberg 9 now finds the numbers in C. virens to be 6 and 3, the lowest numbers 
yet established for plants. The fact that the chromosomes are so few and that 
they are readily recognized as " prochromosomes " in the resting nucleus, removes 
any danger of uncertainty in counting which might be anticipated in case of large 
numbers. Not only is the number low, but the individual chromosomes are not 
alike, two being long, two rather short, and two intermediate. In the diploid 
divisions these three kinds of chromosomes appear in pairs, the members of a 
given pair being alike. At synapsis a double thread appears, and there is a 
fusion which is to be regarded as a fusion of whole chromosomes, reduction in 
number being brought about in this way. Rosenberg suggests that interesting 
results might be obtained by crossing Crepis virens and C. tectorum, and he 
promises to make the attempt. 

Noting the low number of chromosomes in Crepis tectorum (8 and 4) and in 
C. virens (6 and 3), Tahara 10 examined the Japanese species, C. japonica, and 
found the numbers to be 16 and 8, just double the numbers in C. tectorum. The 
chromosomes were also found to be of different sizes and forms. If there is a 
relation between specific characters and chromosomes, the genus Crepis would 
seem to the reviewer to be a favorable form for the investigation of this subject. 

Charles J. Chamberlain. 

9 Rosenberg, O., Zur Kenntniss von den Tetradenteilung der Compositen 

Svensk. Bot. Tidskrift 3:64-77. pi. 1. 1909. 

1° Tahara, M., Ueber die Zahl der C 
Bot. Mag. 24:23-28. pi. 2. 19 10. 



Interaction between scion and stock. — Meyer and Schmidt 11 have produced 
a voluminous article on the interchange of substances and mutual influence between 
stock and scion in a heteroplastic graft. The introduction and review of litera- 
ture occupy 48 pages, and 33 pages are given to the statement of results and the 
summary. One is compelled to think that the article could have been advantage- 
ously condensed to half the space. The authors mention that interchange of 
carbohydrates was already fully worked out, while the previous work on move- 
ment of aplastic and other substances is very unsatisfactory. They direct their 
attention to the movement, formation, and storage of alkaloids, using Nicotiana 
Tabacum as scion on N. affinis and Solatium tuberosum as stocks, and Datura 
Stramonium as scion on Solatium Lycopersicum and 5. tuberosum. They find 
that alkaloids can pass from scion to stock; but the movement is very slow, and 
apparently takes place through the parenchyma and not through the sieve tubes. 
With ^V. Tabacum, normally rich in nicotin, as scion, and N. aftinis, normally 
poor in nicotin, as stock, the latter comes to contain many times its normal amount 
of nicotin, and even ten times as 'much as the scion; while the scion becomes 
relatively poor in it. With 5. tuberosum as stock for N. Tabacum y the periderm 
cells of the former become the main storage tissue for the nicotin. It is most 
abundant in the tissue of the stock just below the graft, and decreases in amount 
as the cells are more distant; while in the tuber none at all or only traces appear. — 

William Crocker. 


Living cells and extreme temperatures. — George vitch, ■ 2 in investigating 
the effect of extreme temperatures on living cells, used the root tips of Galtonia 
candicans. They were kept at 40 C, and — 5 C, killed and fixed at the same 
temperature, and the effect of these extremes noted. At a temperature of +40 C, 



low temperatures the cytoplasm becomes vacuolate, and the coordination in the 
action of the spindle fibers is broken up, which results in the distribution of 
chromosomes between the poles. The activity of the kinoplasm is decreased 
by low temperature and increased by high temperature. There result larger 
spindles with stronger fibers, more rapid transport of chromosomes, and shortened 
duration of nuclear division. Thereby cell wall formation is inhibited and binu- 
cleate cells are of frequent occurrence. The chromosomes often form chains, 

transport toward the poles. In cold preparations the nucleus 


form, also in the warm 

In general it can be said that high temperatures favor development of chromatic 

11 Meyer, Arthur, and Schmidt, Erxst, Ueber die gegenseitige Beeinflussung 
der Symbionten heteroplastischer Transplantationen mit besonderer Beriicksichtigung 
der Wanderung der Alkaloide durch die Pfropfstellen. Flora 100:317-396. figs. J. 

12 George vitch, Peter, Ueber den Einfluss von extremen Temperaturen auf 
die Zellen der Wurzelspitze von Galtonia candicans. Beih. Bot. Centralbl. 25 : 127-135 . 


material, while low temperatures inhibit it. In cold preparations one finds 
collections of chromatin which stain blue and are called pseudonucleoli. 
Schraumex found the same in the cells of shoots of Vicia Faba kept at both high 
and low temperatures. Georgevitch did not find them in warm preparations. 
In cold preparations the nucleoli show an increase in size, mass, and numbers. 

R. Catlin Rose. 

Fossil Osmundaceae. — Kidston and Gwynne-Vatjghan 13 have continued 
their interesting investigations on the fossil Osmundaceae. In the case of the 
most important of the species which they describe (Thamnopteris Schlechtendalii 
Eichwald) there can apparently be no doubt that they have really to do with 
the remains of an osmundaceous fern. They find that in this species the center 
of the stele is marked by the presence of a mass of short tracheids without any 
admixture of parenchyma, which curiously enough they regard as the equivalent 
of a pith. It is surely begging the question as to the origin of medullary struc- 
tures, to regard tissues which admittedly are entirely tracheary and contain not 
the slightest admixture of parenchymatous cells as equivalent to the medulla of 
the higher plants. The difficulty of regarding the central mass of short tracheids 
in Thamnopteris as a pith is rendered insuperable, apparently, by the fact that 
the leaf traces originate from the stele exactly as in those cases where no pith is 
present, that is without giving rise to any foliar gaps. The views entertained by 
the present authors and the majority of English writers on anatomy encounter an 
additional difficulty in that they are quite unable on their hypothesis to explain 
the presence of internal phloem and internal endodermis in closed steles. These 
find apparently a very simple and natural elucidation in connection with the re- 
duction theory now advocated by a considerable number of American anato- 
mists. — E. C. Jeffrey. 

Bennettitales. — Nathorst 14 has described the more or less complete repro- 
ductive apparatus of a number of bennettitean forms. There are three species 
of Williamsonia from the Jurassic beds of Whitby and Scarborough, England. 
In these were found in different cases both microsporangia with microspores, 
and seeds. The structure of the microspores is illustrated by admirable photo- 
micrographs. A new genus (Wielandiella) has a very remarkable vegetative 
organization. The stem branches freely in an apparently dichotomous manner 
and is quite slender. The cones occur in the forkings of the branches. The 
vegetative structure resembles that of the problematic Anomozamites. The 
cones showed remains of both pollen and seeds. The structure of the microspores 
of a third genus (Cycadocephalus Sewardi) is described. These are remarkable 
for their close resemblance to fern spores. For comparison, a figure of Wd- 

J 3 KiDSTON, R., and Gwyxxe-Yaughan, D. T., On the fossil Osmundaceae 
III. Trans. Roy. Soc. Edinburgh 46:1909. 

14 Xathorst, A. G., Paleobotanische Mitteilungen. 8. Handl. Kg!. Svensk 
Vetensk.-Akad. 45: no. 4. 191 o. 


trichia Fr. Braun from the Mesozoic of Franconia is introduced. The result 
of the present important communication is to enlarge our knowledge of the 
male organs of the Bennettitales by seven different species belonging to five dif- 
ferent types. Two species of WUliamsonia have monosporangiate strobili. The 
same condition is clearly demonstrated in Cycadocephalus. The author wisely 




Role of ammonium salts. — Prianischnikow, 15 working with grasses, has 
already shown that a substitution in sand cultures of \-% of the NaN0 3 by (NH 4 ) 2 
S0 4 increases the power of the plant to gain phosphoric acid from raw phos- 
phates (phosphorite), while in absence of (NH 4 ) 2 S0 4 the plants show phosphoric 

acid starvation. 

harvest. Both 

these effects are attributed to the released sulfuric acid. In partial substitution 
the acid was strong enough to aid in dissolving the phosphorite, and in total 
substitution so strong that it greatly injured the plants. It is also shown that 

is very effective in preventing injuries by (NH /t ) 2 S0 4 , and if onlv J-4 

CaC0 3 

enough was used to neutralize the liberated sulfuric acid, the consumption of 
the phosphorite was also much favored. In working with barley, peas, and buck- 
wheat, the author has determined that mixtures of NaN0 3 and (NH 4 ) 2 S0 4 are 
better sources of nitrogen than either one alone, for, as he states, the first is physio- 
logically basic (base liberated due to the consumption of N0 3 as source of nitrogen) 
and the second physiologically acid (acid liberated due to the consumption of 
NH 4 as the source of nitrogen). The two maintain the culture medium neutral. 
The author does not attempt to decide between the relative values of ammonium 
salts and nitrates as a source of nitrogen when the former are of very weak acids, 
as those used by Ritter 16 to settle this question for fungi. — William Crocker. 

Fossil conifers. — Xathorst 17 has described with truly admirable clearness 
and judgment the cones of the problematical coniferous genus Palissya from the 
Rhaetic of Schonen in Sweden. The ovuliferous cone scales are characterized 
by the presence of two rows of opposite seeds, with very loose integuments or 
epimatia. The author concludes that the evidence of the organization of the 
cone scales tends to connect the genus with a second genus described in the article, 
namely Stachytaxus. This genus has yewlike foliage, and attached to the ends 
of the twigs are lax cones with distant scales, each of which bears two ovules, 
provided with a widely flaring integument or possibly an epimatium comparable 
with that found in the Taxineae. The author argues for the taxineous affinities 

J 5 Prianischnikow, D., Zur physiologischen Characterise k der Ammoniumsalze. 
Ber. Deutsch. Bot. Gesell. 262:716-724. 1909. 

16 Ber. Deutsch. Bot. Gesell. 27:582-588. 1909. 

*7 Xathorst, A. G., Paleobotanische Mitteilungen. 7. Handl. Kgl. Svensk. 
Vetensk.-Akad. 43: no. 8. 1909. 


of Stachytaxus, and by implication for the similar relationship of Palissya. 
The only real evidence for the affinity of these two genera with the Taxineae 
seems to rest on the possible presence of an epimatium in connection with the 
seeds. It seems not improbable that they are really representatives of an arau- 
carian stock different from any now in existence. Some of the later mesozoic 
Araucarineae possess both the biovulate cone scale and the flaring integument 
of the genera under discussion. Present indications are that all the mesozoic 
conifers will ultimately be arranged either under the Abietineae or the Araucari- 
neae in the broader sense. — E. C. Jeffrey. 

Orchid flowers and formative stimuli.— Fitting's work on the effect of polli- 
nation and other stimuli upon the postfloration behavior of orchid flowers has 
been reviewed in this journal. 18 In a second paper, 19 he gives an account of 
further experimentation of the same kind, and concludes that the changes induced 
in the perianth, gynostemium, and ovary are at most six, namely: (i) shorten- 
ing of the life of the perianth, (2) lengthening of the life of the perianth, (3) closing 
of the flower, (4) swelling of the ovary and gynostemium, (5) fading of the peri- 
anth, (6) greening of the ovary and perianth. Each of these may result separately 
or with several others, in various combinations. Although it seems probable 
that the influence of the pollen is due to a chemical substance soluble in water 
and alcohol, Fitting was unable to isolate it in pure form or to identify it. It 
was determined, by extracting pollen of Cattleya Trianaei with water and hot 
alcohol, that this chemical substance is not found inside the pollen grain, but 
merely adheres to it and can be removed without injur)' to the pollen. By using 
the pollen from which this substance has been removed, the effects of the pollen 
tube alone can be studied, when it is found that the tube produces the same re- 
sults as the active substance. This is not due to the substance secreted by 
the tube or carried down from the pollen grain, but to an unknown factor. 

R. Catlin Rose. 

A new case of apogamy, — Burmannia coelestis, as described by Ernst, 20 
furnishes a case of apogamy somewhat different from any hitherto reported. 
From the cells of the egg apparatus of an eight-nucleate embryo sac with diploid 
nuclei, one and often two and sometimes three embryos are produced. The 
formation of a tetrad of megaspores is either irregular or completely suppressed, 
as is already known to be the case in most apogamous forms previously described. 
No synapsis stage or heterotypic mitosis was observed. The number of chromo- 
somes was not determined, but is greater than in normally fertilized species of 
Burmannia. The anticipated irregularities in the pollen were found, and the 

i 8 Bot. Gazette 47:479. 1909. 

*9 Fitting, H., Weitere entwickelungsphysiologische Untersuchungen an Orchi- 
deenbliiten. Zeitschr. Bot. 2:225-267. 1910. 

20 Ernst, A., Apogamie bei Burmannia coelestis Don. Ber. Deutsch. Bot 
Gesell. 27:157-168. pi. 7. 1909. 


fact that fertilization is very easily demonstrated in normally fertilized species 
makes the writer confident that the failure to find it in B. coelestis is evidence that 
it does not occur. The figures show only topography, without any details of the 
chromatin situation. A careful counting of chromosomes at critical stages, 
and a few figures at the stages which show whether a form is apogamous or not, 
would have extended the paper but little, and would have made unnecessary 


with these details, the present one being preliminary. — Charles J 

Spermatogenesis in Mnium. — As a result of their studies of several species 
of mosses, the Drs. Van Leeuwen-Reijnvaan reported that in the last division 
of the spermatogenous cells a second numerical reduction of chromosomes takes 
place. In a species of Mnium having eight chromosomes in the last division, two 
long and two short chromosomes pass to the daughter cells. Wilson, 21 studying 
Mnium hornum, in a preliminary note announces that no such reduction is found, 
and that the gametophyte number is constant throughout spermatogenesis. The 
resting nucleus before the final division is quite large and contains a small nucleolus. 
A continuous spirem is not present, and the chromatic material appears as a 
number of small masses from which the chromosomes are formed. In the final 
division the axis coincides with the long axis of the cell, there being no diagonal 



and it is clear that no such reduction as described by the Drs. Van Leeuwen- 
Reijnvaan takes place in Mnium hornum. 

It is to be hoped that the final paper will also deal with fertilization, for many 
investigators find some difficulty in accepting the account given by the Drs. Van 

Leeuwen-Reijnvaan.— W. J. G. Land. 

Hydrogen bacteria.— The epoch-making researches of Winogradski (188 7-) 
on the sulfur, nitrite, and nitrate bacteria established the important fact of the 
existence of non -chlorophyll organisms that are obliged to manufacture their 
organic food by energy obtained from the oxidation of various simple inorganic 
substances. In 1906 various investigators reported the existence of bacteria 





by Winogradski. Lebedefe 22 now makes a preliminary report of the main 
results of an extensive study of the metabolism of these forms. The fixing of 
100 c.c. of C0 2 requires the oxidation of 500-1500 c.c. of H 2 . The oxygen for the 
process is best obtained from atmospheric oxygen, but in absence of it nitrates 
can be decomposed as its source. The oxidation of H 2 still continues in the 
presence of organic food, but no CO. is fixed in that case.— William Crocker. 

21 Wilson, M., Preliminary note on the spermatogenesis of Mnium hornum. 
Annals of Botany 24:235. 191 o. 

22 Lebedeff, A. J., Ueber die Assimilation des Kohlenstoffes bei Wasserstoff 
oxydierenden Bakterien. Ber. Deutsch. Bot. Gesell. 27:598-602. 1910. 


Proteases. — Vines, 23 in continuing his work on the proteolytic enzymes of 
plants, finds that both malt extract and taka-diastase (Parke Davis and Co.) 
contain enzymes capable of digesting fibrin and of splitting peptone. From malt 
extract he has isolated the peptone-splitting enzyme free from the fibrin-digesting 
body, and from taka-diastase he separated each from the other. Both these 
enzymes seem to act best in acid media. In animal tissues there are two fibrin- 
splitting enzymes: a protease, weak and acting in basic media; and /? protease, 
more powerful and acting in acid media. By special methods of preparation, 
Vines obtained a protease which acted best in neutral and basic media. This 
perhaps corresponds to the a protease of animal tissues. The ereptases, peptone- 
splitting enzymes, of animal tissues act best in basic media. Vines's work shows 
that plant ereptases act in acid media. As to terminology, one is inclined to 
believe that Vines could adopt profitably that of animal workers as given by 

Vernon. 24 — William Crocker. 

A new genus of Cordaitales. — Scott and Maslen 25 have described a new 
genus (Mesoxylon) of Cordaitales from the calcareous nodules of the Lower 
Coal-measures of Lancashire. It is intermediate between Poroxylon and Cor- 
daites, as its name implies, including five species which have been referred here- 
tofore to these two genera. The combination of characters is the anatomical 
habit of Cordaites and the centripetal xylem of Poroxylon. The pith is rela- 
tively large and discoid (as in Cordaites) ; the wood is dense, with narrow pith 
rays and relatively small tracheids; the leaf traces are double, but divide before 
entering the leaf; the centripetal xylem is present in the leaf traces at the margin 
of the pith (as in Poroxylon) and throughout their course to the leaves. The 
genus is thought "to completely bridge the gap, so far as anatomy is concerned, 
between the Poroxyleae and the Cordaiteae," and helps to connect the cordaitean 
and later forms (excepting cycadophytes) with the "pteridosperms." — J. M. C. 

Bars of Sanio" in Coniferales — The "bars of Sanio" are "folds" of cellulose 
to be observed in the walls of tracheids as horizontal or more or less semicircular 
markings, which stand out clearly with proper staining. Miss Gerry 26 has 
investigated their distribution among the Coniferales, and has discovered that 
they furnish a constant and useful character in the determination of fossil woods. 
They were found in 35 of the living genera, but do not occur in Agaihis and 
Araucaria, nor in the mesozoic araucarians. Since they do occur in the podo- 
carps, it is concluded that this group is more closely related to the Abietineae than 
to the Araucarineae, a conclusion which contradicts a growing conviction based 


*3 Vines, S. H., Proteases of plants. Annals of Botany 24:213-222. 1010. 

24 Vernon, H. M., Intracellular enzymes. London: John Murray. 1908. 

2 5 Scott, D. H., and Maslen, A. J., On Mesoxylon, a new genus of Cordaitale 
(preliminary note). Annals of Botany 24:236-239. 1910. 

26 Gerry, Eloise, The distribution of the "bars of Sanio" in the Coniferales. 
Annals of Botany 24:119-124. pi. 13. 1910. 




on other characters. It is an interesting fact that the ancient Prepinns shows 
these "bars," which fact helps to establish their ancient character. The total 
result is to emphasize strongly the distinctness of the araucarians from all the 
'other Coniferales. — J. M. C. 

Embryo sacs of some Onagraceae. — An investigation 27 of Epilobium angusti- 
]olium > E. Dodonaei, Oenothera biennis, and Circaea lutetiana shows an interest- 
ing variation from the conventional development of the embryo sac. The usual 
tetrad of four megaspores is formed and the lowest one enlarges and begins to 
develop in the well-known way, but as soon as the four-nucleate stage is reached, 
two synergids and an egg are formed at the micropylar end of the sac, leaving 
one free nucleus in the middle or toward the antipodal end. This sac looks like 
that of Cypripedium, as described by Miss Pace, 28 but is formed from one mega- 
spore, while that of Cypripedium is formed from two. At fertilization, one male 
nucleus fuses with the nucleus of the egg and the other with the single polar 
nucleus, so that there is no triple fusion as in Cypripedium, w r here one of the syner 
gids takes part. The embryo and endosperm develop in the usual way. — Charles 
J. Chamberlaiw 


The original Oenothera Lamarckiana. — Gates 29 has discovered a manu- 
script in the Sturtevant collection of the library of the Missouri Botanical Garden 
" which proves that this plant was originally a species growing wild in Virginia, and 
that it was the first Oenothera introduced into European gardens, about 1614." 
In view of the fact that the origin of this important species has been in doubt, and 
that it has been claimed to have originated in cultivation, this discover)' is note- 



Pinax, by Toannis 

the plant. "The record is as complete and accurate as could be desired, to 
prove to one familiar with the characters of these forms the identity of the plants in 
question." The plant was described under Bauhin's name, Lysimachia lutea 
cormciriata.—J. M. C. 

Color inheritance in Lychnis.— Shull^ has discovered that the purple 
color in L. dioica is a compound character, produced by the interacti<jn of three 
distinct and independent genes. The two types of purple color present in different 
individuals are a reddish purple, changed to blue by alkalies, and a bluish purple, 
changed to red by weak acids. The bluish or alkaline color is hypostatic to the 
reddish or acid color, which is the reverse of the condition found in all other 

2 ? Modilewski, J., Zur Embrvobildung von einigen Onagraceen. Ber. Deutsch. 
Hot. Gesell. 27:287-292. pi i 3 . 1909. 

28 Bot. Gazette 44*.;,53-374- pis. 24-27. 1907. 

29 Gates, R. R., The earliest description of Oenothera Lamarckiana. Science 
S -S. 31:425, 426. I9 i . 

3° Shull, George H., Color inheritance in Lychnis dioica L. Amer. Xat. 44:83- 
9 1 - 1910. 


plants containing similar series of colors. It is inferred that crosses between 
white-flowered plants should result not infrequently in progenies of all purple- 
flowered offspring, or of purple and white in the ratios i : i, 3 : 5, or 1 : 3; but as yet 
these results have not been found. — J. M. C. 

Jurassic flora of Normandy. — Lignier 31 has added a number of new species 
to the rich Jurassic flora of Normandy, that are suggestive of relationships con- 
cerning which real knowledge is very much desired. The Filicales are represented 
by species of Lomatopleris and Linopteris, and the Equisetales by a species of 
Equisetites. The cycadean forms, how r ever, are of chief interest and abundance, 
and it would be a great gain to know definitely what the numerous species of 
Zamites and Otoza mites represent. The conifers are represented by species of 
BrachyphyUum, P achy phyllum , and Coniles. 

The memoir is undated, but its reception in March 19 10 suggests recent 
publication. — J. M. C. 

Apospory and apogamy in Trichomanes. — Georgevitch 32 has investigated 
Trichomanes Kaulfussii, whose apospory and gemma production was described 



(singly or in tufts), at the ends of each of which is balanced a gemma. The 
development of prothallia from these gemmae is described in detail, and antheridia 
were observed developing directly upon the gemmae, sometimes associated with a 
prothallium on the same gemma. This transition from sporophyte to gametophyte 
is accompanied by no reduction in the number of chromosomes. Counts were 
made in both generations and at different stages of mitosis, and always approxi- 
mated 80.— J. M. C. 

Parasitic fungi of Wisconsin. — In 1884 Trelease published a list of the 
parasitic fungi of Wisconsin, and supplementary lists were issued by Davis in 
1893, i ^97j an d 1903. Now a fourth supplementary list has appeared. 33 It 
contains a list of 76 forms occurring on hosts not previously recorded; and 113 
forms not reported heretofore from the state. The latter list includes 9 new 
species and varieties in the following genera: Ascochyia, Cercospora, Cylindro- 
sporium (2), Gloeosporiitm, Phyllosticta (2), Ramularia, and Septoria. This 
record in reference to 189 forms indicates what interest and persistence can do 
for any area. — J. M. C. 

31 Lignier, Octave, Vegetaux fossiles de Normandie. VI. Flore jurassique de 
Mamers (Sarthe). Mem. Soc. Linn. Xormandie 24: pp. 48. pis. 2. figs. 7. (Undated.) 

3 2 Georgevitch, Peter, Preliminary note on apospory and apogamy in Tri- 
chomanes Kauljussii Hk. et Grew. Annals of Botany 24:233, 234. figs. 7. 19*°. 

33 Davis. J. J., Fourth supplementary list of parasitic fungi of Wisconsin. Tran 
Wis. Acad. Sci. 16:739-772. 1909. 


■HH . 



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No. 2 


August 1910 



The Morphology of the Podocarpineae 

Mary S. Young 

The Origin of Ray Tracheids in the Coniferae w. p. Thompson 
On the Relationship between the Length of the Pod and 

J. Arthur Harris 

Fertility and Fecundity in Cercis 


of Zamia 



Oxidizing Enzymes 


Sap Stain 




Briefer Articles 

A Modification 

of a Tung-Thoma Sliding Microtome for Cutting 

R B. Thomson 

Current Literature 

The University of 



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XLbc Botanical (5a3ette 

a /fcontblg Journal Embracing all Departments of JBotanfcal Science 

Edited by John M. Coulter, with the assistance of other members of the botanical staff of the 

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Issued August 18, 1910 



Botanical Laboratory 138 (with plates iv-vi). Mary S. Young 81 


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AND FECUNDITY IN CERCIS (with one figure). J. Arthur Harris - - - 117 


ELORIDANA (with twenty-two figures). Frances Grace Smith * - - - 128 


Irving W. Bailey 142 



FIGURE). R. B. Thomson 148 






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«$s March 



Botanical Gazette 

AUGUST igio 



Mary S. Young 


Previous to 1902 the morphology of the Podocarpineae was an 
unknown field, but recently, through the contributions of Coker (5), 

Jeffrey and Chrysler (7), Burlingame (2), Brooks and Stiles 

(1 . and Young (25) on Podocarpus and Dacrydium, and Noren (i i), 
Stiles (15), Thompson (16, 17, 18), and Tison (22) on Saxego- 
thaea and Mkrocachrys, the group as a whole has become fairly 
well known. Pherosphaera, with two species, is the the only genus as 
yet untouched, and therefore will not be considered in the following 

( 1 cussion. 

hips. The div 



and Pinaceae was made originally on the basis of external 

characters, but with increasing knowledge of the Podoca 

O «•"*"• «,%* & 


mphasized, until it has even been suggested that these two 

hould form a group by themselves. 

Three papers on Phyllodadus by Miss Robertson (13) and Miss 
KUDAHl (8, 9) are of particular interest because of the difference 
of opinion in regard to the affinities of that genus. Phyllodadus was 
c assed with the Podocarpineae by Strasburger, removed to the 
Taxmeae by Engler and Prantl, and finally made a sub-family 


its place is again called in question. 




chiefly anatomical. Miss Kildahl's work was done in this laboratory, 
and I have had the opportunity of examining her preparations. The 



j stages. This work, a previous 
study of Dacrydium, and the opportunity of examining some material 


I shall first 


relationships of the Podocarpineae. 

L Phyllocladus 

The material sent by Dr. L. Cockayne of Christchurch, New- 
Zealand, consisted of staminate cones collected at intervals of a few 
days from October 16 to November 13, and ovulate cones of Novem- 

3, 18, 31, January 8, 28. The iron-alum 

hematoxylin and orange 

very satisfactory 

in staining, as it brought out cytological details remarkably well for 
tissues killed in formalin and alcohol. 


Miss Kildahl reports (1) the formation of two prothallial cells, 
the first of which is usually evanescent; (2) the presence of four, 
occasionally five, free nuclei in the mature pollen grain; and (3) 
the division of the body cell into two equal male cells. 

My material furnishes a close series, beginning with the micro- 

spore stage. The wings are comparatively small and irregular, 
recalling those of Microcachrys, and are formed in the ordinary way 
as cavities in the exine. The intine is usually rather thin. Very 
small starch grains occur, but are never conspicuous. Figs. 1-5 s ^ ^ 
the cutting off of the prothallial cells. The first usually degenerates 
so quickly that it is hard to find in the older grains. In fig. 4 lt lS 
seen partly covered by exine ; this is an oblique section escaping the 
wings and making the spore coats appear uncommonly thick. 

The generative cell is formed in the usual way and divides anti- 
clinally, as in the case of Ginkgo, Podocarpus, and Dacrydium, and 
presumably in all the Podocarpineae. The four-celled stage, as 
shown in fig. 7, but for the wings might very easily be mistaken for 
the shedding stage in Ginkgo. In the latter, it will be remembered* 


the generative cell does not divide until after the tube begins to 
grow. The spindle in Phyllocladus is always more or less oblique, 
and results in the formation of a larger, more centrally placed body 
cell and a small stalk cell. 

In fig. 13 there is shown a peculiar but very common feature of the 
sections, the appearance as of an additional cell cut off from the body 
cell, but without a nucleus. This was noticed also in Podocarpus 
and Dacrydium. Jeffrey and Chrysler (7), in the case of Podo- 
carpus ferruginea and P. dacrydioides, describe a second lateral 
derivative cell and picture it with a nucleus. Brooks and Stiles (i) 
find the same thing in P. spinulosa, but it is not clear whether a 
nucleus is present or not. It is in these species that the most exten- 
sive prothallial tissue is found, consisting of eight cells. In the 
forms studied by Burling ame and myself, as well as in Dacrydium 
and Phyllocladus, no nucleus ever appears in the extra section of 
cytoplasm. The explanation is most easily found in Phyllocladus, 
where horizontal sections show that it is really only a part of the 
stalk cell. When we consider the shape of the generative cell and 
the position of the spindle in the division, it is evident that an oblique 
wall in such a dome-shaped structure could not help resulting in the 
partial encircling of one cell by the other. The situation can be most 
clearly shown by diagrams. A horizontal section in the position 

shown by the dotted line xy in fig. 15 would give a view such as is 
outlined in fig. 16, while fig. 15 is the vertical section through the 
dotted line in fig. 16. Fig. 17 is an oblique section in about the 
position of the dotted line vw in fig. 15. In this case the second 
prothallial cell is seen encircled by the stalk cell. A change of focus 
brings into view the body and tube nuclei and a portion of one wing. 
If the stalk nucleus, as shown in fig. 16, were a little more elongated, 
it would be quite possible to obtain a vertical section in such a position 
as to show a small part of it on each side of the body cell. The appear- 
ance of Jeffrey and Chrysler's and of Brooks and Stiles's 

figures of Podocarpus ferruginea, P. dacrydioides, and P. spinulosa 
suggests this as a possible explanation of the second lateral derivative. 
The fact has been noted that the division of the generative cell is 
anticlinal in Podocarpineae and Phyllocladus, and periclinal in other 
conifers and in cycads, the relative position in the latter having given 


rise to the term " stalk cell." It will be remembered that the division 
takes place in the former before and in the latter after germination 
has begun. Fig. 18 is a diagram showing this stage in Dioon, taken 
from Chamberlain (3). If in the case of Phyllocladus the genera- 
tive cell should enlarge upward toward the tube before the division, 
we can easily see that the oblique wall, failing to touch the prothallial 
cell, would give us the Dioon situation. On the other hand, if the 
wall in Dioon were a little lower down or slightly more oblique, we 
should have a section very like that of Phyllocladus. From this 
point of view the distinction between the anticlinal and periclinal 
division appears to be related to the time at which the division takes 
place, and to be of little significance in itself. 

Throughout the development of the gametophyte, though distinct 
cells are formed, each bounded by a Hautschicht, there is no evidence 
of cellulose walls. This I believe to be true also in the Dacrydium 
and Podocarpus which I have examined. Noren (ii) fails to find 
cellulose walls in the case of Saxegothaea except in the first prothallial 
cell, and Thompson (17) finds them in Microcachrys only in the 
prothallial cells. 

Miss Kildahl mentions the occasional persistence of the first 
prothallial cell. Figs, n and 12 show not only this but also a still 

more rare case in which the second has divided. This, it will be 
remembered, is the usual condition in Dacrydium, Microcachrys, and 
Saxegothaea. In Dacrydium, moreover, the first prothallial cell very 
often degenerates early, which makes the resemblance to Phyllocladus 

still stronger. 

As in all the Podocarpineae so far studied, the prothallial and 
stalk nuclei become free in the general cytoplasm. This may occur 
in Phyllocladus before the grains are shed, but sometimes not until 
after they reach the micropyle. The mature grain contains the bod} 
cell, and the free prothallial, stalk, and tube nuclei. 

Young tubes were found in the nucellus November 20. Fig. l 9 
shows the tube nucleus in advance, followed by the others, and the 
distinctly organized body cell still in the grain. 

My work does not confirm Miss Kildahl's in regard to the nial 
cells, as in every case where a complete series of sections was 
secured, a decided difference in size was evident. The functional 




cell is almost always in advance. It has not only more cytoplasm 
than the other, but nearly always a larger nucleus, and often the other 
shows signs of degeneration. Figs. 20 and 32 are both drawn so as 

to show the largest diameter of each male cell and nucleus. 


division of the cytoplasm is sometimes hard to demonstrate, and the 
appearance is that of two free nuclei. Careful staining, however, 
shows that two definite male cells are formed. 


Some belated cones collected November 18 gave the earliest stages. 
Fig. 21 shows an ovule with its free nucellus, its wide open micropyle 
with pollen grains, and the arillus making its appearance as a slight 
swelling at the base. The stony part of the integument develops 
from a layer two cells deep. The "outer fleshy layer" does not 
thicken up, but remains represented by only the epidermis and the 
cells directlv under it. 


The megaspore mother cell evidently gives rise to a row of three 
cells, the innermost being the functional megaspore; in fig. 3 it 
has germinated and has two free nuclei. The young prothallus is 
surrounded by a layer of glandular, vacuolate cells encroaching on 

the surrounding nucellar tissue. This 


much more marked and the cells become binu 
e membrane soon anoears and is well marked 

at a later 
>ate. The 

free-nucleate stages. 

In having the megaspore membrane and 

spongy layer Phyllocladus agrees with all the Podocarp 
Podocarpus, and differs from all the Taxineae. 

Wall formation was not found, nor archegonium ir 
development of the young gametophy te is evidently rapid, 








of multinucleate cells; Miss Kildahl 




width at the top seems to be due to the rapid centrifugal growth of 
the tissues at the upper (archegonial) end of the prothallus, which is 
indicated by radial lines of cells and numerous spindles. The initial 


has divided, forming a central cell and a primary neck cell; the latter 
gives rise by two divisions to a plate of four cells (figs. 25-27). An 
exceptional case is seen in fig. 29, where a division has occurred in 
each of the four cells. Cellulose walls (in this division), however, are 
not formed, the cells being separated merely by a Hautschicht, and 
in one of them not even a Hautsckicht appears. The cytoplasm of 
the* central cell is very delicate in the early stages, but gradually 
thickens up. The nucleus is rather small and lies at the upper end 

close under the neck. 

There is an interesting peculiarity in the neck development of 
Phyllocladus which, so far as I know, has not been reported in any 
other genus. Miss Kildahl described the pollen tube as pushing 
its way into the archegonium with two apparently detached cells in 
front of it, which she thought were the remains of a crushed neck. 
Fig. 32 shows this stage, but there are four neck cells, and they are 
connected with the adjacent jacket cells by a distinct membrane. 


ery thin and 

from the archegonium in the preparations. 

The origin of this peculiar condition is found in a series of earlier 
stages. The young archegonia are superficial and the necks are 
covered by the heavy megaspore membrane (figs. 24-26). The pollen 
tubes at this time come in contact with the prothallus near the necks, 
and the adjacent tissues grow rapidly, leaving the archegonia at the 
bottom of considerable cavities which are occupied by the tubes 
fig. 36). In Cephalotaxus, according to Coker (4), the archegonia, 
in the absence of pollination, may be entirely inclosed by the growth 
of the lateral tissue. The megaspore membrane is always a little 
thinner at the micropylar end, but disappears entirely inside the 
archegonial cavities (fig. 37), apparently digested by the male gameto- 
phyte. Fig. 27 shows it beginning to disappear under the advancing 
tube. The growth of the female tissues and the pressure of the tube 
result in a lateral stretching, first of the neck, then of the adjoining 
jacket cells (fig. 28). The walls of these latter are able to resist the 
digestive action and enlarge enormously, until just before fertilization 
they appear as the membrane mentioned above (figs. 33-35)- 

An interesting situation is shown in fig. 34. The ovule contained 


three archegonia, of which the two shown had apparently arisen from 
adjacent initials and formed a complex with a common jacket. The 
necks, consisting of four cells each, are attached to one another, one 
on each side of the common wall. This wall is almost parallel with 
the plane of the page and passes between the tw r o egg nuclei. In 
another gametophyte three archegonia were found with a single jacket 
layer separating two of them; the usual amount of tissue lay between 

them and the third. 


action of the tube. The neck of this archegonium has been pushed 
into a vertical position, and part of the egg cytop 


squeezed away from the rest and shows signs of degeneration. 

Shortly before fertilization a ventral canal nucleus is cut off, but 
there is no trace of a wall (figs. 30, 34). In this Phyllocladus agrees 
with Podocarpus, Taxus, and C e phalotaxus , but not with Torreya. 
In the latter not even a nucleus has been found. 

Podocarpus is the only member of its tribe whose female gameto- 
phyte has been studied. In it Coker (5) found six to ten archegonia. 




At the time of fertilization both egg and jacket cells are rich in 
"proteid vacuoles," and the egg nucleus is surrounded by a homoge- 

nous and very dense layer of cytoplasm. The contents of the pollen 
tube enter the egg through the neck, leaving the cells intact. Fig. 
35 shows both male cells in the egg; the larger fuses with the egg 
nucleus, some of the cytoplasm apparently contributing to the embryo. 
Though no trace of the prothallial, stalk, and tube nuclei could be 
found in this case, it is reasonable to suppose that they entered with 
the rest of the contents of the tube. In fig. 27 a tube is seen pressing 
against the side of an archegonium. With further growth probably 
the wall will be forced into a vertical position, in which case fertiliza- 
tion can take place in the usual way. 

Miss Kildahl reports the formation of at least eight free nuclei in 
the proembryo, and my material furnishes nothing more. Ovules 
collected in January had embryos with two cotyledons and long 


suspensors. The entire endosperm tissue is multinucleate, a condi- 
tion which begins to appear while the archegonia are young ; it results 
from the failure of walls to form. In the older endosperm as many 
as eight nuclei occur in a cell. The megaspore membrane at the 
fertilization period is about 2.5^ thick at the lower part of the 
prothallus, and about 4. 5 fi when the cotyledons appear (fig. 31). 

Before discussing relationships, it will be helpful to summarize 
the important facts about Phyllocladus, including the work of Miss 
Robertson and Miss Kildahl. 


1. The stamen bears two abaxial sporangia. 

2. The pollen grains have two wings. 

3. In the male gametophyte there are two prothallial cells; the 
first usually disappears, the second occasionally divides. 

4. The generative cell divides anticlinally but obliquely, and the 
stalk cell partially encircles the second prothallial and body cells. 

5. The mature pollen grain contains the body cell and the free 
stalk, prothallial, and tube nuclei. 


6. There are two unequal male cells, only the larger of ^ 

7. The ovulate structure is a strobilus with a single erect ov 
in the axil of each scale. 

8. A symmetrical arillus originates from the base of the ovule and 
remains free from the integument. 

9. The outer fleshy layer is represented by two layers of cells. 

10. The nucellus is free to the base. 

11. The megaspore mother cell givt 


the innermost of which is the functioning megaspore. 

12. The spongy layer and megaspore membrane are strongl} 

13. Each ovule contains two archegonia, occasionally three or 
four, each with its own jacket, but sometimes there is a complex 
two in a common jacket. 



occasionally these divide anticlinally. 
15. There is a ventral canal nucleus. 


16. The mature archegonium is very wide at the top and is 

covered by a membrane formed from the walls of the adjacent jacket 

17. In fertilization the contents of the pollen tube pass through the 
neck, leaving it intact. 

18. Some male cytoplasm contributes to the embryo. 

19. There are at least eight free nuclei in the proembryo. 

20. The embryo has two cotyledons and a very long suspensor. 

21. Mesarch bundles occur in the cladodes. 

22. Taxinean sculpturing is found in the tracheids. 

23. No vascular strands enter the ovule; the ovular supply con- 
sists of two strands facing each other and ending in a tracheal plate 
below the integument. 


The characters which point toward the affinity of Phyllocladus 
with the Taxineae are: (1) the structure of the ovule, (2) the sym- 
metrical arillus, (3) resemblances of the ovulate cone to that of 
Cephalotaxus, (4) mesarch bundles, and (5) taxinean sculpturing of 
tracheids. Characters indicating a relationship with the Podo- 
carpineae are: (1) the character of the male gametophyte, (2) the 
structure of the stamen, (3) winged pollen grains, (4) megaspore 
membrane and spongy tissue. 

The first point is perhaps the strongest argument for taxad affinity, 
as the erect, free, axillary ovule is characteristic of the whole group. 
This ovule, however, is a primitive type which we would expect to 


ase of any line. Though the progress of the 
ard inversion and fusion of parts, which finds 
1 Podocarpus itself, we find many relativel) 

stages represented. In Dacrydium, Saxegothaea, and Microcachrys 
the ovule is free from the scale and epimatium, except at the base, 
and in the various species of Dacrydium we find all positions from 
erect to completely inverted. The young ovule of Saxegothaea is in 
early stages perpendicular to the scale and becomes inverted only as 
the result of later growth. This, together with its position near the 
base of the scale, is suggestive of ancestors with erect, axillary ovules. 
The homology of the symmetrical arillus which originates from 


the base of the ovule, and of the one-sided epimatium arising from 
the scale is uncertain. The origin of the latter may be related, how- 
ever, to the foliar origin of the ovules, and, as Miss Robertson states, 
" the asymmetry is correlated with the inverted position of the ovule, 
so that it will not do to lay too much stress on this point, as proving 
that epimatium and arillus are not homologous." 

Though the external resemblance between the cones of Phyllo- 
cladus and Cephalotaxus is rather striking and the vascular supply is 
the same, this may merely point back to a common origin of all the 
Taxineae. Cephalotaxus, moreover, has two ovules in the axil of 
each bract, while Phyllocladus agrees with the Podocarpineae in 
having only one. 

The presence of mesarch wood in the cladodes owes its significance 


any other group of conifers. This, too, is an ancestral character, for 

members of anv group. Stiles 

finds it in the ovular supply in Saxegothaea, whose cone is regarded as 
the most primitive of the podocarps. 


if it were confined to Taxineae, but this is by no means true. Such 
characters as these last two are valuable in connection with other 
evidence, but have little weight in themselves. 

The most convincing evidence of podocarp affinity lies in the 
entire behavior of the male gametophyte : the formation of prothallial 
tissue, the freeing of the nuclei, and the early division of the genera- 
tive cell. The presence or absence of prothallial cells is a definite 



all. Permanent prothallial cells are known nowhere in conifers 
except in Podocarpineae and Araucarineae ; and the early division 
of the generative cell, which characterizes the former, occurs no- 
where else except perhaps in araucarians. Prothallial tissue is a 
primitive character, possessed, as we presume, by the ancestors of all . 
conifers; but the Taxineae have eliminated it entirely, while it still 
remains one of the most characteristic features of the Podocarpineae. 
The morphology of the different forms of stamen in the conifers 
is an open question. The Taxineae, according to Coulter and 

1 9 1 o] YO UNG—PODOCA RPI XEA E g i 


family, except araucarians, has a stamen with more than two pollen 
sacs. If the two types have a common origin, only the Taxus stamen 
can be the primitive one; and in this case Phyllocladus has gone a 
long way in the podocarp line of development. A different origin, of 


Winged pollen is entirely absent among the Taxineae, but is 
characteristic of the Pcdocarpineae, being absent only in the 
Saxegothaea. The irregularity in size and 


Microcachrys has given rise to Thompson's (16, 17) theory that the 
two-winged condition has developed within the group and shows no 
relation to Abietineae. This gives us another podocarp line along 




It is entirely eliminated in Taxineae, 


carpus itself. As the ovulate structures of the type genus are by far 
the most specialized of the family, this is not surprising. Miss 
Robertson thought the megaspore membrane and spongy tissue 
were in some way correlated with the presence of winged pollen, but 
their occurrence in Saxegothaea with its wingless microspores breaks 
down this supposition. 



the whole that: (1) Phyllocladus has primitive characters of the 
Taxineae which are being eliminated in the Podocarpineae ; (2) it 
has primitive characters of the Podocarpineae which have been 
entirely eliminated in the Taxineae; (3) it has some advanced char- 
acters of Podocarpineae ; (4) the taxad resemblances are on the whole 
more superficial and variable, and the podocarp features more 
fundamental; (5) the resemblances to Podocarpineae are too strong 
to justify the retention of the intermediate family. 

We conclude, therefore, that Phyllocladus is a relatively primitive 
member of the Podocarpineae, which branched off from them a 
comparatively short time after their separation from Taxineae. 




II. Podocarpineae and Araucarineae 

The more important contrasts between the araucarian and podo- 
carp lines are in the structure of the stamen and ovulate strobilus, 
and in the method of fertilization. The araucarian stamen is a 
comparatively primitive type and is somewhat suggestive of Ginkgo. 
It is one-sided and bears three to thirty pendent pollen sacs; while 
all the genera of the Podocarpineae, as before mentioned, have the 
microsporangia definitely reduced to tw r o. 

The ovulate cone in Araucarineae is a compact structure with 
many scales, ripening dry; while the podocarp line is characterized 
by the reduction in the number and size of cone scales and the tendency 
to fleshy development. Saxegothaea, the form which stands nearest 
the araucarians, shows the least amount of reduction in both size 
and number of scales; Microcachrys is next; while in other forms 
the cone is represented by the single, apparently terminal, ovule and a 
few rudimentary scales. The podocarp ovules have the arillus, or 
epimatium, which is absent in araucarians. The cone scale of 
Araucaria, on the other hand, bears the so-called ligule, represented 
in Agathis by only a slight projection from the surface. The ovules 
of Saxegothaea are united with the scale only at the base, while in 
Araucaria they are described as imbedded in the tissues of the scale. 
In Agathis, however, the ovules are free and the seeds winged. 

The female gametophyte is too little known in either group for 
any adequate comparison. That of the Araucarineae, however, is 
apparently much more primitive than that of the known podocarps. 
The archegonia are very numerous and are described (Seward and 
Ford 14) as situated at the bottom of deep pits and usually not 
connected with the surface by necks. Investigation is needed here 
to show whether they are really hypodermal, or whether the condition 
is brought about by overgrowth of adjacent tissue in the development 
of a neck so massive as not to have been recognized. The position of 
the ovule of Araucarineae in the tissues of the scale is another subject 
which needs interpretation. 

Fertilization in Araucarineae (Thompson 19) is angiosperm-like, 




The first suggestion of relationship between the two groups came 
from the study of the male gametophyte. This has recently been 
supplemented by studies of Saxegothaea, which have brought out 
striking resemblances to certain species of araucarians. The geo- 
graphic distribution is also suggestive of alliance between these two 
great southern groups. 


In the Araucarineae, as in the Podocarpineae, there are two 
original prothallial cells, from which by subsequent divisions a more 
or less extensive tissue is formed. In Araucarineae there may be 
as many as thirty cells (Lopriore 10; Thompson 21). Jeffrey 
and Chrysler (7) found an apparently mature pollen grain of 
Agathis australis with a prothallial complex of eight cells, which 
is the situation in some species of Podocarpus. Other species of 
Podocarpus have four cells, resulting from a single division of each 
original prothallial cell. In Dacrydium, Saxegothaea, and Micro- 
cachrys there are two to four permanent prothallial cells, in Phyllo- 
cladus one to three. Thus we have a complete overlapping prothallial 
reduction series from Agathis to Phyllocladus. 

The generative cell is found in Araucarineae as in the others, but 
the division into stalk and body cell has not been observed. Unless 
this division takes place in the tube, as is true of most conifers, it 
must either have been eliminated or missed in the preparations. The 
otherwise close correspondence with the gametophyte of Podocar- 
pmeae inclines one toward the latter supposition. Further work is 
needed on this point. 

the ovule and female gametophyte 
The single inverted ovule is characteristic of both families. The 


inversion was probably developed independently, however, in the 
two lines. In Saxegothaea, as has been said before, the young ovule 
changes from an erect to an inverted position in the course of its 
development The free nucellus of Dacrydium and Phyllocladus is 

• , m 

a primitive feature which they have in common with araucarians. 
The female gametophyte, as before stated, is little known. A 


point of resemblance, however, between Araucarineae, Phyllocladus, 

and Podocarpus is fc 
in the archegonium. 



The resemblance of Saxegothaea to the araucarians, noted by 

Stiles (15), Noren (ii), Thompson (18), and Tison (22), consists 

chiefly of anatomical characters and certain external features of the 
cones. Stiles finds in the stem tracheids somewhat araucarian in 
character, " an occasional tendency to a two-ranked arrangement of 
the pits, and in these cases the pits become alternate and hexagonal. 
Two-ranked and alternate pits horizontally flattened have also been 
found in Dacrydium. The prominence of transfusion tissue is another 
character suggestive of araucarian affinity. 


The staminate cone is described by Stiles as somewhat araucarian 
in general appearance, and the w r all of the microsporangium as strik- 
ingly like that of Araucaria Rulei. Brooks and Stiles (i), in their 
study of Podocarpus spinulosa, state that the wall of the sporangium 
is very like that of Saxegothaea and Araucaria. Wingless pollen 
is another point of contact. The fact that the pollen grains of all 
the other Podocarpineae have wings does not affect the argument, 
if we accept Thompson's theory that they were developed within the 
group (15, 16). 

In the ovulate cones of both Saxegothaea and Araucaria, especially 
A. Rulei, there is a gradual transition from foliage leaves to sporo- 
phylls. This and the similarity in the vascular anatomy are con- 
sidered by Thompson and Tison as indications of the simple nature 
of the strobilus in these groups. The occurrence of a single resin 
duct in the sporophjll is, according to Stiles, another indication 01 
this in Saxegothaea. 

One of the most striking features of Saxegothaea is the projection 
of the nucellar tissue through the micropyle, where it expands to 
form a stigma-like knob. The same thing occurs less conspicuous!) 
in Araucarineae, but, with the exception of a few abnormal cases 1 
angiosperms, is elsewhere unknown. This feature appears to 
related physiologically to the difficulty of fertilization, and its mor- 
phological significance is doubtful. Thompson (19) sees in u 



tendency toward protosiphonogamic fertilization, significant in relat- 
ing Saxegothaea to the araucarians. 

A good deal of stress has been laid recently on the distribution of 
the vascular bundles in the ovulate sporophylls. Stiles finds the 
branching in Saxegothaea very similar in the main to Araucaria 
Rulei and A. Cookei, and the arrangement in Microcachrys, though 
not unlike that of Saxegothaea, resembles more closely A . Bidwillii. 
Tison agrees essentially with Stiles in regard to the details in Saxe- 
gothaea, but finds greater resemblance to A. brasiliana and A. imbri- 




and the simple structure of the ovulate scale in the two families, but 
they differ somewhat in the application of their results. Stiles 



ales are descended from some common ancestor with its micro 

and megasporophylls both arranged spirally in cones. Along one 
line of descent we find the Araucarieae, along the other is Saxego- 
thaea leading on to Microcachrys and the other Podocarpeae." 
Noren comes to the same conclusion, regarding Saxegothaea as 


ocarpineae ; 

more closely related 

latter group, and regards Microcachrys as the connecting link. 

But it is the attempt to homologize the ovulate structures of conifers 
that has given rise to the greatest amount of discussion. In conifers 



9 » * 

5«s its supply being inverse to the other. The two systems may 


also i 

from the other. The former varies 

from a considerably 



made a study of the distribution of the 



very complete historical account of the controversy on the morphology 
of the ovulate cone. He agrees with Celakovsky that the arillus 
of Taxaceae, the ligule of Araucaria, and the ovuliferous scale of 
the other Pinaceae are homologous and are all a second integument, 
that all the ovulate cones are morphologically compound, but that the 
sporophyll is suppressed and represented by the ovule alone. 

Thompson (18) in 1909 brought out some interesting data in regard 
to the inversion and its relation to the theory of the axillary shoot. 
The inversion of the bundles supplying the ovule he explains as 
normal for sporangial supply, and cites cases of such inversion in the 
microsporophylls of cycads and of some conifers. In Tsuga he found 
two inversions, the ovular supply being inverse to that of the scale, 
which in turn is inverse to that of the bract. The first inversion he 
considers as homologous with that of the scale of Saxegothaea and 

related to the ovule, while the second may be explained by the theory 
of the compound nature of the sporophyll. On this basis he makes 
two groups of conifers, the Araucarineae and Podocarpineae having 
simple strobili and ovules on the morphological upper surface of the 
scale, and the other Pinaceae having compound strobili and ovules 
morphologically abaxial. 

Tison regards the arillus of Saxegothaea and the ligule of Ara il- 
ea ria as homologous with the ovuliferous scale of other conifers, 
calling them an ovuliferous appendage. He does not commit 
himself in regard to the axillary shoot theory as affecting Abietineae, 
Cupressineae, and Taxodineae, but agrees with Thompson that the 
cones of Saxegothaea, Podocarpineae, and Araucarineae are simple- 
He favors the inclusion of Saxegothaea and Podocarpineae in the 
Araucariales suggested by Seward and Ford (14). 


After reviewing the whole situation, one is impressed with the 


that they are probably related, but that the question is by no means 
settled. The whole coniter group still appears as a maze of cross 
resemblances. If we confine ourselves to one or two characters, 
the problems of relationship are comparatively simple, but bring 
more into consideration, and we are immediately in trouble. The 


relation of the podocarps with Araucarineae on the one hand, for 
instance, is complicated with evidence of connection through Phyllo- 
cladus with taxads on the other; and the two last-named tribes are 
apparently very widely separated from one another. It is true that 
the evidence for the first relation is the stronger, for without Pkyllo- 
cladus the question of taxad affinity would hardly be raised at all; 
while without Saxegothaea we should still have much evidence of the 
other connection. The existence of Phyllodadus, however, cannot 



conclusiveness. In the first place, it is based largely on primitive 
characters. These may indicate merely that neither group has 
advanced far from the original ancestral conifer stock. The two 
lines may be quite distinct and both short. Two short branches from 


of the same long branch. Most 



cular bundles. Such evidence is unsatisfactory on account of the 
great variability of the structures concerned. Great variations are 
found between closely related forms. Araucaria with its ligule and 
apparently imbedded ovule, and Agathis with its free winged seed, 
or the ovulate structures of different species of Dacrydium, are ex- 
amples. Tison finds variations in the arrai 


bundles of Saxegothaea, not only in different individuals, but in the 
same plant. He says: "En ce qui concerne les ecailles fertiles, je 
dois tout d'abord faire remarquer que la disposition du systeme 
fasciculaire a leur base, dans la region ovulifere, est tres variable 
souvent dans un meme cdne, ces variantes n'etant pas necessaire- 
ment en rapport avec la position des bractees sur les cones." 




simDle. and one is inclined to doubt the mor 

Phological need of such homologies, 
lines of resemblance amon 


g conifers noint to a more 

for all the families than we are apt to think, and in this case the sig- 



The greatest gap in our present knowledge of conifers is shown 
by the Araucarineae. Even in the male gametophyte, which supplies 
perhaps the strongest argument in the above discussion, a good deal 
of work is needed. The female gametophyte is little know T n and 
the embryo practically not at all. Beside this, we should know the 
early development of the ovulate strobilus for a proper understanding 
of the morphology of its parts. In the Podocarpineae we lack 
adequate knowledge of the female gametophyte, embryo, and develop- 
ment of ovulate structures. Until further data on these points are 
available, we should be hardly justified in coming to a definite deci- 
sion in regard to relationships, and at present it seems best to hold 
Taxineae, Podocarpineae, and Araucarineae apart as separate tribes, 
leaving open the question of larger groupings among conifers. 

Grateful acknowledgments are due to Professors J. M. Coulter 
and C. J. Chamberlain, under whose direction this work was done. 

The University of Chicago 



i. Brooks, F. T., and Stiles, W., The structure of Podocarpus spin 
(Smith) R. Br. Annals of Botany 24:305-318. pi. 21. 19 10. 

2. Burlingame, L. L., The staminate cone and male gametophyte of Podo- 
carpus. Bot. Gazette 46:161-178. pis. 8, g. 1908. 

3. Chamberlain, C J., Spermatogenesis in Dioon edule. Bot. Gazette 

47-215-236. pi. 15- 1909- 
4. Coker, W. C, Fertilization and embryogeny in Cephalotaxus ForUinei. 

Bot. Gazette 43:1-10. pi. 1. 1907. 

5- , Notes on the gametophytes and embryo of Podocarpus. Bot. 

Gazetie 33:89-107. pis. 5-7. 1902. 

6. Coulter, J. M., and Land, W. J. G., Gametophytes and embryo of Torreya 
taxijolia. Bot. Gazette 39:161-178. pis. A, i-j. 1905. 

7. Jeffrey, E. C, and Chrysler, M. A., The microgametophyte of the Podo- 
carpineae. Amer. Nat. 41:355-364. 1907. 

8. Kildahl, N. J., The morphology of Phyll 
46:339-347. pis. 20-22. 1908. 

9» , Affinities of Phyllocladus. Bot. ( 

0. Lopriore, G., Ueber die Vielkem 

schlauche von Araucaria Bidwillii. Ber. Deutsch. Bot. Gesell. 23 :335'3-* 6 

pi. 15. 1905. 

Bot. Gazette 



11. Noren, C. O., Zur Kenntniss der Entwickelung von Saxegothaea con- 
spicua Lindl. Svensk. Botanisk. Tidskrift 2:101-122. pis. 7-9. 1908. 

12. Pilger, R., Taxaceae in Das Pflanzenreich (Engler) 4:5. 1903. 

13. Robertson, A., Some points in the morphology of Phyllocladus alpina 
Hook. Annals of Botany 20:259-265. pis. 17, 18. 1906. 

14. Seward, A. C, and Ford, S. O., The Araucarieae, recent and extinct. 
Phil. Trans. Roy. Soc. London B 198:305-411. pis. 23, 24. 1906. 

15. Stiles, W., The anatomy of Saxegothaea conspicua Lindl. New Phytologist 
7:209-222. 1908. 

16. Thompson, R. B., Note on the pollen of Microcachrys. Bot. Gazette 
46:465. 1908. 

l 7> , On the pollen of Microcachrys tetragona. Bot. Gazette 47:26-29. 

pis. j, 2. 1909. 

J 8. — , The megasporophylls of Saxegothaea and Microcachrys. Bot. 

Gazette 47-345-354. pis. 22-25. 1 


l 9> , The Araucarieae; a protosiphonogamic method of fertilization. 

Science N.S. 25:271, 272. 1907. 
20. - ? The megaspore membrane of the gymnos perms. Univ. Toronto 

Studies, Biol. Ser. no. 4. pis. 1-5. 1905. 
2I - , Preliminary note on the Araucarineae. Science N.S. 22:85. 1905. 

22. Tison, A., Sur le Saxegothaea conspicua Lindl. Mem. Soc. Linn. Nor- 
mandie 23:139-160. pis. 9, 10. 1909. 

23. Worsoell, W. C, Observations on the vascular anatomy of the female 
"flowers" of Coniferae. Annals of Botany 13:527-548. pi. 27. 1899. 

2 4- , The structure of the female "flowers" in Coniferae. Annals of 

Botany 14:39-82. 1900. 
25. Young, M. S., The male gametophyte of Dacrydium. Bot. Gazette 
44=189-196. pi. IQ. 1907. 


t, tV, and tV mm. were used. All drawings, except figs. 15, 16, and 18, were 
made with the aid of the camera lucida. The magnifications refer to the figures 
as they appear on the plates, after having been reduced one-half in reproduction. 

Abbreviations: p ly first prothallial cell; p 2 , second prothallial cell; g, genera- 
tive cell; s, stalk cell; b, body cell; /, tube nucleus; a, arillus; n, neck; m t , 
functioning male cell; «,, functionless male cell; vn, ventral canal nucleus; 


FlG. i— Microspore. X950. 
FlG. 2.— Spindle for the first prothallial cell. X950. 
Fig. 3.— First prothallial cell cut off. X950. 
• IG. 4— Spindle for the second prothallial cell; fin 
cove red by inline. X950. 



Fig. 5. — Second prothallial cell cut off, the first degenerating. X950. 

Fig. 6.— Spindle for generative cell. X950. 

Fig. 7.— Tube nucleus, generative cell, and prothallial cells. X950. 

Figs. 8, 9. — Generative cell dividing. X950. 

Fig. 10. — Mature pollen grain. X950. 

Figs, ii, 12. — First prothallial cell persistent and the second divided. X950. 

Fig. 13. — Mature grain, showing a part of the stalk cell on each side of the 


body cell. X950. 

Fig. 14. — Nuclei becoming free. X950. 

Fig. 15. — Diagram of vertical section through dotted line in fig. 16. 

Fig. 16. — Diagram of horizontal section through dotted line xy of fig- i5> 
showing form of stalk cell. 

Fig. 17. — Horizontal section, showing stalk and second prothallial cells and 
tube and body nuclei. X950. 

Fig. 18. — Outline drawing of Dioon. — From Chamberlain. 

Fig. 19. — Body cell in the grain; stalk, tube, and prothallial nuclei in the 
tube. X440. 

Fig.»2o. — Body cell divided, forming two unequal male cells. X44°- 

Fig. 21. — Longitudinal section of ovule and bract; pollen grains on the 
nucellus. X45. 

Fig. 22. — Young stage of the arillus; detail of fig. 21. X44°- 

Fig. 23. — Two-nucleate stage of female gametophyte; detail of fig. 21. X44°- 

Fig. 24. — Young archegonium, showing primary neck and central cells. 



Fig. 25. — Young archegonium; primary neck cell divided. X44°- 

Fig. 26. — A neck cell dividing. X440. 

Fig. 27. — Four-celled neck; megaspore membrane disappearing before 
advancing pollen tube; the space between the membrane and neck cells is due 
to shrinkage. X440. 

Fig. 28. — Stretching of adjacent jacket cells under pressure of the pollen 
tube; body cell and central nucleus not yet divided. X440. 

Fig. 29.— Eight-celled neck. X440. 

Fig. 30. — Egg and ventral canal nucleus. X440. 

Fig. 31. — Endosperm cells and megaspore membrane; January 28. X95°- 

Fig. 32.— Egg ready for fertilization; four-celled neck and jacket membrane, 
male cells, stalk, prothallial, and tube nuclei. X440. 

Fig. 33.— Effect of pollen tube on an archegonium; the membrane intact. 


Fig. 34. — Two archegonia in a common jacket;^ two pollen tubes. X34 a 

Fig. 35. — Fertilization. X440. 

Fig. 36. — Female gametophyte. X45. 

Fig. 37. — Megaspore membrane and archegonial cavities. X44°- 



>N S\Uvx*v 

} 7H XG on Pin ZLOGLi U)f 5 



> Vf WG on P/n 7 J, OCL M)l S 



YOUNG on P/mXOCL. V)f S 



W. P. Thompson 

(with SIXTEEN figures) 


phylogenetic significance of ray tracheids in the Coniferae. Pen 


from their distribution and, from, his theory 


ui ineir origin, concludes that the number in a given species is in 
direct proportion to its specialization, and tKat the forms where they 
are most numerous are derived from those where they are not so well 
developed. Jeffrey (2), on the other hand, from a study of certain 
traumatic phenomena, considers that the forms where they attain 
their greatest development, namely the pines, are the most ancestral, 
and have given rise by degeneration to the ones in which they occur 
sporadically. Apart from its intrinsic interest, it is hoped that the 
present study, by determining the origin of the ray tracheid, will 
supply a basis for its correct phylogenetic interpretation. 

In carrying on the study, a thorough investigation was made of 
the character and mode of formation of the ray tracheid throughout 
the individual plant, but more especially in the primitive regions: 
seedling stem and root, young branch and young root of the adult, 
and the axis of the seed cone. The 

forms . 

cone. The forms chosen for detail 
igenous species of the hard and soft pii 
>bus, but the results were confirmed in n 
method was to follow a rav by means < 

Pin us 




origin of the medullary ray at the pith has been described and 
by Kny (3) for Pinus silvestris. He states that in this region 
all the ray cells are parenchymatous, and elongated not radially but 
vertically. These long cells are often in connection with similar cells 
from rays lying above or below. 


Very soon they s 
direction, and elongate radially to form 


A later 






writer, Conwentz (4), agrees with this description and extends it 
to include the root. I have found the observations of both writers 
to be practically true for Pinus Strobus and P. resinosa. In view 
of what is to follow, however, dissent must be expressed from the 
statement that during their further course the rays are separate. An 
important feature which neither writer emphasizes is that the rays 



essential differences; accordingly these regions are treated separately. 


Some distance from the pith certain peculiar tr ache ids appear, 
which Kny and other investigators seem to have overlooked. 

Fig. 1 

Fig. 2 

Figs, i, 2. — Fig. 1, P. resinosa: young root; short tracheids extending between 
two rays; fig. 2, P. Strobus: young root; short tracheids with characteristic radial 

Examples of them are shown in figs, i and 2. They occur in radial 
rows, extending between two rays which lie in the same vertical 
plane. Consequently they are of various lengths, depending on the 
distance between the rays. Their form is also variable, often being 

Quite uniform, like tracheids (fijr. tV often vprv iVrpcnilar in outline. 


A frequent peculiarity is the possession of arms (fig. 2) projecting 
radially either from the ends, or the middle, or from both, and meet- 
ing similar projections from their fellows. In this case the bodies 
of the cells are some distance apart. The walls of all these short 

tracheids are generally thinner, and their bordered pits much smaller 
than those of ordinary tracheids. 

The usual sequence of these cells and their transformations 
toward the cambium are illustrated in fig. 3, which is a camera lucida 
drawing of tw r o consecutive sections from a series of the root of Pinus 
resinosa. The cambium in this figure is toward the right (this 
orientation has been preserved throughout my illustrations). On 
their first appearance, toward the left of the figure, these tracheids 
have the more regular form approximating ordinary tracheids. 
Farther out they become irregular, usually with the above mentioned 
projections; then divisions occur, forming tw r o superposed rows. 
These soon separate, taking up their position along the rays (center 
of figure) and forming irregular marginal cells, which gradually 
become regular and assume the form of true ray tracheids. In the 
ray at the bottom of the figure, rav tracheids have already been 
formed, but in one at the top this process has not yet been completed. 
In Pinus Strobus the sequence is similar. Often in both, the series is 
not so regular as the one just described; some stages may be hurried 
over, some greatly prolonged. In the one drawn, the transformation 
has taken place much more quickly than is usually the case, for the 
transitional stages may often be traced through several years' growth. 

That these elements are indeed transitional is further shown in 
% 3 (Pinus resinosa) by the development on their walls of the denta- 
tions characteristic of the ray tracheids of the hard pines. When 
they first appear the elements are quite smooth, but as they assume 
more the shape and character of ray tracheids they acquire these 
dentations. One may often find intermediate cells whose walls are 
partly smooth like tracheids and partly dentate like ray tracheids 
(center of fig. 3) . 

It is evident that in the root of these forms we have a complete 
transition between short tracheids and ray tracheids; that by a 
process of division, of shortening, and of radial extension marginal 
ray tracheids have been produced from tracheary tissue. 





The description so far has dealt with the marginal ray tracheids. 
The interspersed ones appear about the same time; their origin is 
illustrated in fig. 4. At the left of the figure are the irregular short 
tracheids extending between two rays. Farther to the left, that is, 
nearer the pith, these rays are more distant and the transitional 
tracheids are longer. To the right the rays gradually approach and 

Fig. 3. — P. resinosa: young root; the transformation of short tracheids to 
marginal ray tracheids. 

the tracheids shorten to form the regular interspersed ray tracheids 

which continue to the cambium. 



ones. In the latter case the rays separate and the tracheids form 
along their margin. In the former, the rays draw together and the 
tracheids shorten and take their place in the center of the resulting 
composite ray. 

The formation of this ray calls attention to an undescribed phenome- 
non apparently rather common in Pinus, namely the fusion of rays. 
As noted above, when the rays originate at the pith they are usually 


In the root wood 
made to determin 




• — o 

It was found that 





ery one of the high rays was formed 



:e lower ones. Only very rarely was a height of four 
otherwise. The so-called primary rays are then really 
formed secondarily, by fusion. To what extent this fusion is char- 
acteristic of the other forms has not yet been determined, but it would 
seem that the low ray is the primitive condition in the pines. 



Figs. 4, 5. — Fig. 4, P. resinosa: young root; the transformation of short tracheids 
to interspersed ray tracheids, and the fusion of rays; fig. 5, P. Strobus: young root; 
the replacement of ray tracheids by parenchyma; at x is a degenerating tracheid. 

The approach of the rays does not always end in the production of 

interspersed tracheids 



on the point of doing so, then these elements are often replaced by a 
row of parenchyma cells which soon assume the ordinary form and 
size. In fig. 5 at the left are the short transitional tracheids which are 

cells. Farther toward the 


cambium these gradually become lower and indistinguishable from 
other cells of the ray. 1 Instances were also observed where the 

* In this drawing, and in the others as well, with the exception of tigs. 8, 9, 14-16, 
the simple pitting of the wall of the parenchyma has been used to distinguish this tissue 
rom Ae tr acheary elements with their bordered pits, the nuclear and protoplasmic 
contents of the former being omitted. 




fusion of rays eliminated a row of tracheids without replacement by 
parenchyma. The tracheids were simply "pinched" out, only the 
parenchyma cells continuing. In a few cases, even after true marginal 
ray tracheids had been formed as above indicated, the rays drew 
together and fused, with the result that the marginal ray tracheids 
were "pinched" out. 





Figs. 6, 7. — Fig. 6, P. resinosa: young root; illustrating the formation of 
double row of ray tracheids; fig. 7, P. resinosa: young root; formation of a wholly 
tracheidal ray, and its transformation into a parenchymatous one. 

The origin of the marginal and interspersed ray tracheids and 
their relationship to the fusion of rays have been described. The 
more complicated case of the formation of two rows of tracheids on 
the margin of a ray is partly illustrated in fig. 6. At the left, toward 
the pith, are two rows of transitional tracheids touching end to end. 
Beyond the figure, nearer the medulla, these have been formed from a 
single row of longer tracheids. Toward the cambium the row on the 




margin of the upper ray settles down to form regular ray tracheids 
while the other remains transitional. A repetition of the process in 
this row gives rise, beyond the figure, to a double series on the margin 
of the upper ray. In many cases, however, the second row r is formed 
without the intervention of the transitional cells, this stage being 
either hurried over or completely omitted. 

The only kind of ray tracheid whose formation has not yet been 
described is that composing the wholly tracheidal ray. When the 
rays are far apart, two or more divisions may take place in the long 
tracheids instead of the usual single one. Then one or more rows of 
transitional cells are formed midway between the rays, with other 
rows above and below, that is, touching the rays. When the latter 
rows separate from the central ones, these continue as irregular 
tracheidal cells which gradually become regular, forming a com- 
pletely tracheidal ray. Fig. 7 represents only the ends of the series, 
a considerable space at the center being left out; in a are two rows 
of irregular tracheids, the lower of which settles down to form the 
completely tracheidal ray seen in b. 

At the right in figure 76 another phenomenon is illustrated. This 
is the replacement of a tracheidal ray by a parenchymatous one. It 
is not an abnormality in this section, for 


numerous examples were seen. In fact 
it seems to be the common method of 



to the pith. Owing to 
the yearly increase in the circumference 
of the wood, many new parenchyma rays 
must be formed if the number in a given 
area is to remain at all constant, and it 
seems easier for them to be produced 
from cambial cells which give rise to ray 
tracheids than from longer ones which 

w r ood 

Fig. 8.— P. resinosa: young 
root; the development of a 
ray tracheid at the cambium. 


cambium further evidence is afforded 

at the right. 

structures from tracheary 

In fig. 8 the cambium is 






"turning" along the ray and developing the buttresses characteristic 
of the ray tracheids of the hard pines. The lower part remains 
smooth. At this stage then, it is quite intermediate between a \ 
tracheid and a ray tracheid. 

Farther from the cambium whole rows of tracheidal cells may be 
observed, each with a long tail-like projection extending from one 



111 * I 111 

Figs. 9, 10. — Fig. g, P. resinosa: young root; a row of ray tracheids with tail-like 

projections; fig. 10, P. Strobus: adult root; ray tracheids with " tails. 

end (fig. 9) , and this end is always the one nearer the cambium. The 
"tail" is evidently the smooth part of the tracheid as seen at the 
cambium. Moreover, in Finns resinosa the "tail" lacks the char- 
acteristic dentations present on the rest of the cell. Apparently then 
it is the result of an incomplete shortening of the tracheid. Fig. i°» 
from an old root of Finns Strobus, shows that these projections persist 
in the mature wood, but, as is to be expected, they are neither so 
numerous nor so conspicuous. 


admittedly more consen 


lonary processes 






series can rarely be observed. Nevertheless, the transitional form of 
tracheid is occasionally found, and has been drawn in fig. 1 1 from the 
second year's growth of Pinus Strobus. Tail-like projections are 
quite common, and have been illustrated in t 

J 11 




11, 12. 

-Fig. 11, P. Strobus: young stem; transitional tracheids; fig. 12 
P. Strobus: adult stem; ray tracheids with "tails." 

The only good stem series showing the transformation from 
tracheids to ray tracheids was observed in old stem wood of Pinus 
resinosa which had been wounded (fig. 13). The rapidly shortening 
series of tracheids ends at the right in true ray tracheids with buttressed 
w alls. In this wounded material also the tail-like projections were 
v ery numerous and very large. Both features are to be accounted 
for as traumatic reversions similar to those described by Jeffrey for 
Cunninghamia (2), in which form though ray tracheids are not 
normally present they were found in connection with a wound. 

Even in the normal adult ray tracheid, however, we find indica- 
tions of its tracheary origin. One of these is the occurrence of tertiary 




spirals. This has been described by Bailey (5) for some species of 
Pinus as wall as for Picea and Pseudotsuga. He states: "Ray 
tracheids appear to follow closely the wood tracheids. Where spirals 

Fig. 13. — P. resinosa: adult stem; a series of tracheids from a wounded region, 
showing transformation to ray tracheids. 

are strongly developed (summer wood) in the latter elements, the} 
will also appear in the former." This concomitant occurrence 01 
tertiary spirals on the elements in question argues community 01 

• * 



So far the study of the seedling has presented no features not 
observed in the corresponding regions of the adult, except that here 

the ray tracheids are later in appearing. 

Cone axis 

True ray tracheids are quite absent from the cone axis, as Jeffr 
and Chrysler (6) have pointed out. I have found their place taken, 
however, by the bent ends of the tracheids. Fig. 14 shows a trachei 




whose end is bent along a ray for a remarkable distance. Such 

tracheids are very common in the 

cone axis, more than half the rays 

showing at least one in some part of 

their short course. Evidently they do 

the work of the ray tracheid, and 

probably represent the first step in a 

process of turning and shortening 
which, as described above, ends in 
the production of ray tracheids. 
Similar bent ends are sometimes to 
be found in the young stem, and are 
evidently to be interpreted in the sa 

Fig. 14. — P. Strobus: cone axis: the 
end of a tracheid bent along a ray. 

Relation to albuminous cells 

In the light of what has been said of the origin of ray tracheids, 
an interesting circumstance is furnished by their relationship to the 
albuminous cells of the ray. Strasburger (7) points out the 
homology between the latter and sieve tubes. "The albuminous 
cells are higher than the ordinary cells of the ray and like sieve tubes 
have sieve areas, soon lose their contents, and ultimately collapse." 
They are then virtually ray sieve tubes. Now in the Abietineae, 
where ray tracheids are numerous, albuminous cells were always 
found conterminous with them through the cambial region. This is 
illustrated in fig. 15 even for a row of interspersed ones. In the Arau- 
canneae and Taxaceae, where there are no ray tracheids, there are no 
albuminous cells. In those Taxodineae and Cupressineae where no 
ray tracheids were seen, no albuminous cells were found. And in 
Thuja, where ray tracheids occur sporadically, the only albuminous 
cells were those lining up with ray tracheids. Albuminous cells are 
then always associated with ray tracheids (an interesting exception 
is noted below). This association and their sieve tube character 
indicate that they bear the same relationship to ray tracheids as the 
sieve tubes to wood tracheids. They thus afford valuable collateral 
evidence of the tracheary origin of their representatives in the wood. 

The exception referred to is found in Abies balsamea. Here almost 
every ray passing through the bast has albuminous cells on its margin. 






line with the parenchyma cells of the ray, but always above or below 
them. Often they are in line with two or three degenerating cells on 
the wood side. These facts support the view of Jeffrey (2) that 

the scarcitv of rav tracheids in certain Abietineae. including Abies. 

is the result of degenera- 
tion. The albuminous 
cells, elsewhere insepa- 
rable companions of the 
ray tracheids, persist, 
while the ray tracheids 
themselves have disap- 
peared. The latter are 


Fig. 15. — P. resinosa: old stem; cambial'region 
showing albuminous cells in line with ray tracheids. 

probably repre 

by the degenerating cells 


Another observation 

supporting this view is 
that of the occurrence 
of ray tracheids in the 

wounded root of Abies amabilis. Hitherto all observers have reported 
ray tracheids absent from the wood of this species. Yet in a piece 
of root wood which had been wounded several times, undoubted ray 
tracheids were developed in considerable numbers. This phenomenon 
parallels the traumatic revival of ray tracheids in Cunningham^ 
described by Jeffrey (2), and admits of a similar interpretation, m 
this case their ancestral presence in the genus Abies. 

General considerations 

and relationshios of the rav tracheid 



As has been 

shown, it is in the young root that the proof of its origin from tracheary 
tissue is most conclusive. Here transitions w r ere observed from shor 
tracheids extending between the rays to ray tracheids of all EXPOS 

with these transitional areas, is the occurrence of degenerating eel s. 
One of these has been incidentally illustrated in fig. 5 at *. I he ^ 




are structureless, shadowy outlines replacing transitional tracheids, 
which they resemble in form, although often somewhat more irregular. 
They occur in considerable numbers wherever the former elements 
are found, and are evidently to be regarded as "degeneration prod- 

ucts, to which transitional structures are always subject. As such 
they emphasize the transitional character of the regions in question, 
and help to complete the chain of evidence for the origin of the ray 
tracheids from tracheary tissue. 

Penhallow's argument (1) for his theory that ray tracheids are 
derived by modification from the parenchymatous cells of the ray, is 
the occurrence of ray 
tracheids conterminous 
with parenchyma cells. 
But this is so rare that 
DeBary (8) was led to 
assert that it never 
occurred. I must con- 
firm Penhallow's ob- 
servation, however, Fig. 16.— P. serotitia: medullary ray showing (1) 

especially in the young the ray tracheids with dentate walls, (2) the structure 

plant, where such an 

appearance is more 

of the parenchyma cells, (3) tracheids conterminous 
with parenchyma cells. — From Pexhallow (i). 

common than in the adult. I have observed and figured this feature 
m the formation of secondary parenchymatous rays from wholly 
tracheidal ones (fig. 7), and in the replacement of interspersed ray 
tracheids by parenchyma (fig. 5). I have observed also a similar 
replacement of marginal ray tracheids by parenchyma cells. In these 



Miix ^^ lliv , , 1&U1 , „* mv. " /7 - - 

tracheary elements, the reverse of what is required from Penhallow's 

point of view. 

indicated in the very figure which Pexhallow 



The direction in 

which the oblique end walls are inclined, and in which the tail-like 
Projection points, which is seen on one of the ray tracheids on the top 
of the ray, indicates, as shown above (figs. 8-10, 12), that the cambium 

in i. . . I 1 . _ - * 1 • 




with the ray tracheid is then nearer the cambium, and so what we 
really have is the replacement of a row of tracheary elements by 
parenchymatous ones. The origin of such a row of ray tracheids is 
to be looked for toward the pith, and, as I have shown above, the ray 
tracheids arise here either in connection with transitional tracheids. 
or, when these are omitted, above or below the ray and not in line 
with parenchyma cells. 

The intimate study of the medullary ray from its beginning at 
the pith in the different regions of the individual plant, besides dis 
closing the origin of the various types of ray tracheid, has drawn 
attention to important features in the distribution of these structures. 
Kny has called attention to their absence in the first year's growth 
of the stem, and Jeffrey and Chrysler state that they are not present 
in the seed cone. In the young branch of Finns Strobus they appear 
sporadically during the second and third years, and then increase 
slowly in number until the adult condition is attained after about ten 
years. In the seedling they are still later in appearing, as is also the 
case in the young root. In the latter region they never become so 
numerous as in the stem. In Pinus resinosa they appear in abun- 
dance much earlier and increase in number much more rapidly, reach- 
ing the adult condition in five or six years. They are never found in 
the cone axis of either species. Thus in the primitive regions of the 
plant ray tracheids do not occur, and therefore they must be regarded 
as specializations. That they are of cenogenetic origin is further indi- 
cated by the fact that in the older pines, the Pityoxyla of the Creta- 
ceous, as described by Jeffrey and Chrysler (6), no ray tracheid- 

In view of their origin and distribution, ray tracheids are regarded 
as specialized structures and their phylogenetic meaning so inter- 
preted. Those woods in which they are most abundant are con- 
sidered most modern, unless, as in Abies, it can be shown that they 
have been secondarily lost. Their character when present is as 
important as their number. For example, if the early growth shou=> 
many transitional elements or a large development of tail-like pro- 
jections, then the wood is stamped as primitive. Again, the smootn- 
wall form in the soft pine is more tracheid-like and therefore more 
primitive than the dentate form of the hard pine, an inference wo* 


is strengthened by the earlier and more rapid development of ray 
tracheids in Pinus resinosa than in Pinus Strobus. 



complete transitions may 



j ^ * vvw ^w**-j^*^i,^ uuujxuviio xxxtxjr ks\, vu^m.u xxwaxx 

short tracheids extending between the rays, to ray tracheids both 
marginal and interspersed. In the young stem only remnants of the 
transition usually remain. The complete series, however, may 
occur traumatically. 

2. Further evidence of the origin of ray tracheids from tracheary 
tissue is found in (i) their develo] 
young plant, (2) the occurrence < 
possession of tertiary spirals. 

3. The occurrence of ray tracheids bears a definite relation to that 
of albuminous cells. 

4. In Abies the possession of albuminous cells and the traumatic 
occurrence of ray tracheids indicate that the latter are vestigial. 

5. The regional and fossil distribution of ray tracheids indicates 
their ancestral absence in the pines. 

6. The hard pines are more specialized than soft ones. 

7- The large rays of Pinus are usually formed by the fusion of 
smaller ones. 

8. Ray tracheids are often replaced by parenchyma cells. The 
importance of this in the formation of secondary parenchymatous 
rays has been indicated. 

ggestion of Mr 

son and carried on with his constant advice. My warmest thanks 
are due to him for quite exceptional kindness throughout the course 
of the work. 

University of Toronto 

1. Pen hallo 




Jeffrey, E. C, Traumatic ray tracheids in Cunninghamia sinensis. Annals 


of Botany 22:593-602. pL 31. 1908. 

Kn "V, L. ? Anatomie des Holzes von Pinus silvestris. 



4. Conwentz, H., Monographic der Baltischen Bernsteinbaume. Danzig. 

5- Bailey, I. W., The structure of the wood in the Pineae. Bot. Gazette 

48:47-55- PL 5- *9°9- 

6. Jeffrey, E. C, and Chrysler, M. A., On cretaceous Pityoxyla. Bot. 

Gazette 42 : 1-15. 1906. 

7. Strasburger, E., Lehrbuch der Botanik. Jena. 1908. 

8. DeBary, A., Comparative anatomy of phanerogams and ferns. English 
edition. Oxford. 1884. 




J. Arthur Harris 

(with ONE figure) 

It is well known to botanists that the egg cell is not the only struc- 
ture affected by fertilization. Goebel points out 1 that in the Hepati- 
cae an accompanying result is often seen in the production of a further 
development of the envelopes of the ripening sporangium. Again, 
in the same work he suggests 2 that the stimulus exercised by pollina- 
tion in the flowering plants is most probably chemical. The prepara- 
tion of the ovules for fertilization is dependent in some cases upon 
pollination. For instance, in such plants as Corylus, Alnus, Quercus, 
and some of their allies, there is no sign of the placenta in the ovary, 
to say nothing of ovules, at the time of pollination. In most species 
of Orchidaceae the ovules are laid down at the time of pollination, 
but still are rudimentary. The stimulus exercised by the pollen tube 
induces the further development of the female sexual apparatus in 
these plants. 

Pfeffer 3 joins with Goebel in considering that the penetration 
of the pollen tube may serve as a stimulus to the development of the 
ovary, and cites the seedless fruits studied by Muller-Thurgau 
and the observations on Ficus by Treub. 

Jost* writes : " The germination of the pollen tube has an exciting 
influence on the development of the fruit. This is particularly 
noticeable in certain cultivated plants, which, as for example currants 
and sultana raisins, produce no seeds, the ovules having degenerated. 
If the stigmas of these plants be not pollinated, the fruit fails to 
develop, but pollination leads to development without leading to any 

1 Organography of plants 2:105. 1902. 

2 Op. cit. 1:269-270. 1900. 

3 Physiology of plants (English transl.) 2:173. 1900-1906. 


u 7 ] 

o. 1907. 


1 1 8 BOTA XICA L GAZETTE [august 

This is not the place for a review of the literature bearing upon 
these interesting chemical, physiological, and morphogenetic prob- 
lems, however one wishes to designate them. The views of the three 
botanists just cited are sufficient to show the interest which is being 
taken in these problems. Several writers have contributed to the 
literature. Without any attempt at arrangement for priority or 
extent of investigation, I mention Ewart (1,2), Solacolu (7), Treub 

(8), Muller-Thurgau (5), Noll (6), and Fitting (3). Fitting 

(4) has recently given a review, with a bibliography, of the chief 
literature in connection with an account of his own work on the 
Orchidaceae. From his experimental studies in this family he is 
led to the conclusion that the stimuli involved in the ontogeny of 
the fruit are in part due to an organic substance, not an enzyme, 
external to the pollen grain, in part to the growth of the pollen tubes, 
and finally in part to the development of the fertilized ovules. 

The observations of the authors mentioned above deal chiefly 
with the influence of pollination as a stimulus inducing the develop- 
ment of the ovary up to a stage where the fertilization of the ovules 
is possible, or a little beyond. These phases of the problem are 
much more easily studied experimentally than that of the influence 
of the developing ovule upon the growth of the ovary. The present 
investigation bears upon this point. 

If the developing seed excretes some substance which acts as a 
stimulus to the development of the ovary wall, or in some other manner 
exerts an influence upon it. it seems not unreasonable to suppose that 
the effect would be greater if several seeds were developing than 11 
there w r ere only one or a very few. 

Our probltm is essentially this: Does the number of ovules which 
develop to matured seeds influence the size of the fruit, and to what 
extent ? 

In all of the pods which have matured at least one seed, the stimulus 
to development due solely to the penetration of the pollen tube (as 
distinguished from the possible influence of the developing zygote) 
should be the same for all pods, unless the quantity of pollen tubes 
which penetrate the tissue of the style differs from ovary to ovary, 
and the intensity of the stimulus is to some extent proportional to 
their number. The problem is surrounded by a good many difficul- 


ties. There seem 

between the number of seeds developing and the size of the fruit 
might arise. 

a. Fruits with a larger number of ovules are apt to be larger, if 
for no other reason, simply because the placental space required is 
greater. In the lone run fruits with larger numbers of ovules also 


have larger numbers of seeds, 5 and an influence on fruit length at 
first attributed directly to the development of the seeds might be due 
indirectly to the number of ovules. 

b. The space required for the matured seeds might, through the 
purely mechanical effects of crowding, result in the fruits with greater 


numbers of seeds being larger in size. 

c The developing seed might by means of some excreted product, 
or in some manner not yet suggested, directly induce a greater develop- 
ment of the ovary. 

d. Both the number of ovules developing into seeds and the size 
attained by the fruit may be to some measure dependent upon some 
other character; say, for example, the position of the fruit on the 
inflorescence axis, or the number of fruits developing per inflorescence. 
The correlation between them might then be due merely to their 
mutual dependence upon some other character. 

In the literature one finds only a few references to the relationship 
between the fertility of a fruit and its size. Ewart (i) gives tables 
showing numbers of seeds and mean weight of fruit for three series 
of I2 5> 48, and 134 fruits of one variety of apples, which indicate 
that weight increases with* number of seeds. According to Ewart, 
Muller-Thurgau found that in apples and pears the size of the fruit 
and the number of seeds are interdependent. Muller-Thurgau 
cut off four of the five stigmas in the pear blossom, fertilized the re- 
maining one, and thus produced asymmetrical fruit. Ewart con- 
cludes : " Es steht demnach ohne zweif el f est, dass den Kern einen 
Wachtumsreiz auf die zur Fruchtbildung bestimmten Gewebe 



In view of the four possibilities suggested above, I think it is 
quite evident that great caution should be used in asserting that the 
number of ovules developing into seeds has per se any influence upon 

5 This statement is based on the results of many series of unpublished observations. 




the size of the fruit. I believe that in such cases the analytical 
methods of modern higher statistics applied to large bodies of data 
are fitted to give results of real value. 

The purpose of the present paper is to present the results of an 
attempt to measure the intensity of the interrelationship between the 
length of the pod and (a) the number of ovules formed, (b) the number 
of seeds developing, and (c) the fecundity, that is, the ratio of number 
of seeds developing to number of ovules formed per pod, in Ceras 

As material for a first study, the number of ovules formed and the 
number of seeds developing per pod were counted and the length 
measured in 3,000 pods, collected at Meramec Highlands, St. Louis, 
Missouri, in the autumn of 1905. To secure as representative 
material as possible, 50 pods from each of 60 individuals were taken. 
The measurements of pod length were taken to the nearest milli- 


poses of calculation. 

Fertility and length of pod for 3,000 Cercis fruits 


• • » 

J • • ■ • 
3 • • * * 









• - 

4 • * * * 

• - 

■ • 

« • ■ 



• * 

• • 

■ • 

• ♦ 


« * 



















• * 














































147 255 

* * 





• • * 


















• t 

■ * 













• * • 


















t • t 















J 3 




























* ■ 


• * 

» • 

# ■ 

• • 

• - 



















Table I gives the data. In the first column to the left the number 
of ovules formed and the number of seeds developing per pod are 


shown in the form of fractions, in which the number of ovules is 
given as the denominator and the number of seeds developing as 
the numerator. For each of the twenty-four seed-ovule classes found 
in our material, the frequencies of the different pod lengths are tabu- 
lated out. 

The physical constants describing these characters are : 

Average ovules 4.6947±o.oni 

Standard deviation of ovules o .9041 ±0 .0079 

Coefficient of variation of ovules 19 . 258 

Average seeds 40613:^0.0134 

Standard deviation of seeds 1 . 0849 ± o . 0094 

Coefficient of variation of seeds 26 . 712 

Average length of pod 76 . i8i±o .095 

Standard deviation of length of pod 6 7 . 68i4±o .0669 

Coefficient of variation of length of pod 10 .496 

Average index, seeds/ovules o. 8650 ±0.00 20 

Standard deviation of index o. i587±o.ooi4 

These constants require no discussion here; they enable us to 
pass to the determination of the degree of interdependence of the 
fertility characters and the length of the fruit. 

Consider first the correlation between number of ovules per pod 
and the length of the pod. We find r ol = 0.4278 + 0.0101. Remem- 
bering that correlation is measured on a scale of o to ± i , we see (a) 
that the sign of the relationship is positive, that is, that as the number 
of ovules per pod increases the length also increases, and (b) that the 
relationship is a moderately close one. 

The degree of interdependence between the two characters may 
be made clearer by expressing it in terms of regression instead of 
correlation. The equation to the regression straight line is 

^ = 56.119 + 3.634^. 

In this case y= length of pod and x— number of ovules per pod. 
From this equation we see that a pod having an ovule more than 
the average of the population would be 3.6 mm. longer than the 
average length. 

6 Sheppard's correction was applied to the second moment in the calculation of 
the constants for length of pod. It was not used for the fertility characters, where we 
are dealing with integral variates, nor in the seed-ovule indices. 






i a Gram 

To the 


















the observations are relatively few. We may 

6 - 

Mean length of pod in millimeters 








t. iIps and ^ 

Fig. i. — Diagram of slope of regression: straight lines, length on ovuiea 

seeds; solid line, theoretical length of pod for ovules; dotted line, theoretical eng 

pod for seeds; solid dots, observed length for ovules; circles, observed average 

for seeds. 


the amount of discrepancy between the theoretical line and einpin 
means by taking the average deviation on both sides of the line gi 
by the equation. Considering the weight to be attached to a a& ia 
to be proportional to the number of observations on which it is bas 



mm. A d 


seem to most biologists quite negligible, but I rather suspect that 


cant. The 


For number of seeds maturing per pod and length of pod we 
find the interdependence ^ = 0.5055 ±0.0092, and the equation to 
the regression straight line 

^ = 58.645 + 3-579^ 

: significance of y is as above, and x= seeds per pod. The 
[fig. 1) shows the agreement between the line given by the 
and the empirical means. Throughout the central region 

gllWLiL HH. ^-^-'""" --» 


good, but at both ends the observed means fall considerably below 
those to be expected from the equation. If these deviations are 
biologically significant, and not due merely to the small number of 
observations which fall at the extremes of the range, they indicate 
that both the pods developing only a single seed and those developing 
the maximum number of seeds are somewhat dwarfed in length as 
compared with the whole series of 3,000 pods. I am not yet ready 
to discuss the reasons for this condition. 

we find 


For ovules and length, r=o.4278±o.oioi 
For seeds and length, r =o.5o55±o .0092 

Difference=o .0777 

Since the correlation is actually, though only slightly, higher for 
seeds developing than it is for number of ovules formed, it would 
appear that the number of seeds developing must have some connec- 
tion with the length of the pod independent of the interdependence 
f °r length and ovules. 

As pointed out early in this paper, one of the difficulties in assert- 
in g that there is a real physiological relationship between the number 
°f seeds developing and the size of the fruit arises from the fact that 




the correlation between the number of ovules formed and the size 
of the fruit may be of such a magnitude that it is impossible to tell 
without relatively refined statistical analysis whether the influence 
apparently due directly to the seeds may not be referred to the ovules. 
In the present case, for instance/the correlation for number of seeds 
developing exceeds only very slightly that for number of ovules formed. 
We may determine whether the number of seeds developing has 
per se any correlation with the length of the pod by the following 
simple process. We sort our material into classes according to the 

number of ovules per 



for each of these subgroups. If now the length of the pod and the 
number of seeds developing are in some measure interdependent, we 
should expect to find significantly different mean lengths for pods ot 
the same number of ovules, but different numbers of matured seeds. 


Mean length of pods for different numbers of seeds per pod in the four 

chief ovule classes 

of seeds 


Number of ovules per fod 









79 7 6 

Such an arrangement of the data is given in table II for the classes 
in which the number of observations is sufficiently large to give smooth 
results. Here the results are very clearly favorable to the hypothesis 
of a relationship between the number of seeds developing and the 
length of the pod, independent of the relationship for number 
ovules, for the mean increases as the number of seeds increases when 





has the advantage of giving a terse quantitative statement of t e 
independent relationship between the number of seeds develops 


and the length of the pod. The partial correlation coefficient gives 
the correlation between two characters, say seeds (s) and length (/) 
for constant values of a third character, number of ovules (0). 

A necessary preliminary is to determine the correlation between 
number of ovules per pod and number of seeds developing per pod. 
We find r os =o. 7297 ±0.0058. Clearly with such a large value 
for the correlation between ovules and seeds we would expect some 
relationship between the number of seeds developing and the length 
of the pod, having no direct physiological significance whatever, but 
due merely to the fact that since number of ovules and number of 
seeds are closely correlated, and number of ovules and length of pod 
are correlated, number of seeds and length of pod must also be corre- 
lated. It is the influence of the ovules which we wish to remove by 
means of the partial correlation coefficient. The familiar formula 



''si ^os^ol 

1 i-r 2 /i-r 2 , 

which gives 


Psl = 0.3I28±O.OIII. 7 

I think this is a rather significant result. It not only shows that 
there is a physiological or morphogenetic relationship between the 
number of seeds developing and the length of the fruit independent 
of the correlation for ovules and length, but tells us the intensity of 
the interdependence as well. 

There is still another way in which the influence- of the ovules may 
be, to some extent at least, cleared away. Instead of correlating 

maturing per pod and 



ratio or index seeds/ovules per pod) and 
be found: fa-o.2906io.0113. This 
constant indicates very clearly that there is a real interdependence 
of number of seeds developing and fruit length, which is independent 
of the correlation for number of ovules and length of fruit. 

7 The probable error of p s l is from the formula £^[ = 0.67440. 1 -p*\/\/n. Mr. 
D AVm Heron, of University College, London, tells me that he has recently demon- 
strated the correctness of this formula and has the proof in press. 


Comparing the two methods of obtaining the independent corre- 
lation between number of seeds and length we have : 

Correlation of index and length o . 29o6±o .0113 

Partial correlation coefficient o.3i28±o.oin 

Difference 0.0222 

This difference is certainly of no practical importance. 


1. In considering the influence of the number of seeds develop- 
ing upon the dimensions attained by the fruit, the number of ovules 
formed cannot be disregarded, since a correlation attributed directly 
to the influence of the development of the seeds may be in part at least 
due to an interdependence between the number of ovules formed and 
the dimensions of the fruit. The influence of the number of ovules 
can be neglected only when the coefficient of correlation between 


number of ovules and size of fruit is demonstrated to be zero. 1 m* 
point is well illustrated by a series of 3,000 pods of Cercis, where the 
correlations for ovules and length is r o[ = 0.428 ±0.010; while the 

correlation for seeds per pod and length is only r sl = o . 506 ± o . 009. 

2. Two methods for freeing the correlation between the number 
of seeds developing and the length of the fruit from the influence 
the relationship between the number of ovules formed and the leng 
of the pod are suggested; the first is the determination of the partia 
correlation coefficient, that is, the correlation between number 

seeds and length for constant values of numbers of ovules per pod- 
the second is the determination of the correlation between the inae* 

Seeds developing pe r pod , , t , - - , 

"O^Tes formed per pod" an( * the length of the pod. 

The results from the data in hand are in close agreement 



PsI— - ril = 0.022 

and we conclude that of the gross correlation of about p = ° co ° 



say P* 


ship between the number of seeds developing and the length 01 




1. Ewart, R., Blutenbiologie und Tragbarkeit unserer Obstbaume. Land- 


Jahrb. 35: 1906 

Berlin. 1907. 


3. Fitting, H., Die Beeinflussung der Orchideenblii ten durch die Bestaubung 

und durch andere Umstande. Zeitschr. Botanik. 1 : 1909. 
4- •, Entwickelungsphysiologische Probleme der Fruchtbildung. Biol. 

Central bl. 29:193-206, 225-239. 1909. 
5. Muller-Thurgau, H., Kernlose Traubenbeeren und Obstfruchte. Land- 

wirtsch. Jahrb. Schweiz. 1908. 
0. Moll, F., Ueber Fruchtbildung ohne vorausgegangene Bestaubung (Par- 

thenocarpie) bie der Gurke. Sitzungsber. Niederrh. Geo. Nat. Heilkunde. 



7. Solacolu, T., Sur les fruits parthenocarpiques. Compt. Rend. Acad. Sci 
Paris 141: 1905. 

8. Treub, M., L'action des tubes polliniques sur le developpement des ovules 
chez les Orchidees. Ann. Jard. Bot. Buitenzorg. 3: 1883. 



Frances Grace Smith 

(with twenty-two figures) 

The results of a study of the development of the staminate strobilus 
and microsporangia of Zamia were published by the writer in 1907 (1). 
Some of the material, sent from Miami, Florida, in June of the years 
1905 and 1906 for this investigation, included ovulate cones. Later 
in the year 1906 an effort was made to secure a complete series 01 
young ovulate cones, for Zamia, alone of the cycads of North America 
exists in such profusion that whole plants may be sacrificed to secure 
a single small cone from each. There are some stages not yet covered 
by the series of cones which has been obtained, but it seems worm 
while at this point in the study of the material to sum up the results, 
and to postpone conclusions from these results and their theoretical 
bearing upon other cycad studies until a complete series has been 

Each year, since 1906, an attempt has been made to secure materia 
which should give the origin of the integument. In 1907, out 


eight or ten plants sent from Florida between July 25 and August 
not one contained an ovulate cone. I do not know whether this was 
an unfruitful year or whether the collector was unfortunate in the 
plants he gathered. Another year, knowing just the period durin}- 
which the material ought to be gathered, careful collections were made, 
but in every case the cones had reached a development two weeKS 
ahead of that of the previous year, so that it is evident collections 
Zamia must be made often and during long periods in order to obtai 
a full series. 

The facts ascertained from the material cover the period from t 
appearance of the ovulate cone to the time when the developing 
endosperm has partially filled the embryo sac, and will be trea e 
under three periods of growth. 

Botanical Gazette, vol. 50! 






i. Development of the strobilus and the sporophylls 

Plants gathered early in June show a slight elevation which elon- 
gates to form the strobilus, but at this stage it is impossible to dis- 
tinguish the ovulate strobili from the staminate. As stated in my 
description of the staminate 
plants (1), the strobili are 
deeply sunken in the tip of 
the crown and are completely 
covered by the bases of the 
rosette of leaves. 

A strobilus of July 5 is 
about 5 mm. long, and at this 
time can be recognized as 
ovulate. There are defined 
in a median longitudinal sec- 
tion five sporophylls on a 
side (fig. 1). A comparison 
with a staminate cone of 
July 8, with eleven sporo- 
phylls on a side, gives ap- 




but greater breadth and larger 
sporophylls than those of the 
staminate cone (fig. 2). 

In comparing cross- Figs, i, 2— Fig. 1, longitudinal section of 

1 e young ovulate strobilus, showing sporophylls 

: n y 2 > ( j u i y 5 ). x 4 o; % 2, longitudinal section of 

"men the same points are yoU ng staminate strobilus, showing sporophylls 

noticed. A staminate (Julys). X40. 

strobilus (fig. 3) has on an 

average fourteen to sixteen sporophylls on a cross-section, while the 

ovulate strobilus (fig.' 4) has seldom more than ten. A difference in 

the size of the sporophylls as well as of the strobili is seen also. 

The development of the sporophylls is identical in the two strobili, 
at least in these early stages, except that the elevation representing the 

sporophyll in the ovulate strobilu: * "" 

"i a hypodermal position which 
sporangia appear, there are fewer 






sporophylls, as shown in figs. 3 and 4. The lobes themselves are 
broader and the cells of the inner surface show a deeper stain, indi- 
cating that this is the region where active growth is taking place. 
This region is included by the dotted line drawn on one of the sporo- 
phylls in fig. 4. 

In a megasporophyll of July 25, of the size in fig. 4, a cell can be 
distinguished for the first time from those about it by its size and 

Figs. 3, 4.— Fig. 3, transverse section of staminate strobilus, showing spo 
phyllsand position of sporangia (July 26). X40; fig. 4, transverse section of ovua 
strobilus (July 26), showing sporophylls and meristematic group of cells on one 
indicated by dotted line. X 40. 

by its larger nucleus with deeply staining chromatin. This eel 


hypodermal in origin (fig. 5) and resembles the single archespo 
cell in the staminate sporangium. About ten or twelve cells m 
cross-section form a meristematic group, some of which are actney 
dividing, but this archesporial cell is easily distinguished from these. 
In another sporophyll of July 25, but evidently one a little m° r 

developed as shown by its increased size, 

the development of this 

group of cells has gone on still farther, .and now four cells m crok 
section can be clearly distinguished from the others (fig- °)- 
group is separated from the epidermis by one layer of cells, and 




size of its nuclei and the position of its walls suggest an origin from a 
single archesporial cell. It resembles, too, the group in the staminate 
strobilus which gives rise to the sporogenous mass in the microspo- 
rangium. Two pairs of cells (I) just under the epidermis, from the 

il of sporophyll of fig. 4, showing hypodermal arche- 
e advanced stage, showing group of meristematic cells 

Ros. 5r 6.— Fig. 5, detail of sporoph, „. „ 

pona cell. X40; fig. 6, more advanced stage, showing group 
\ > which may give rise to integument. X 1400. 


isions of hypodermal 



ation of the integument. 

referred to, and the strobili of August 8, which is the date of the next 
collection, show on either lobe of the sporophyll the projecting nucellus 
and the tegument, which is only slightly elevated at this time. 




2. Development of the ovule 

The ovule of August 8 shown in figs. 7 and 8 is younger, I think, 
than that of Stangeria as figured by Lang (2), and about the same 
age as that of Ceratozamia shown by Treub (3). The megasporv 
mother cell is easily picked out at this stage as a larger cell, which is 

Figs. 7, 8. 



cells arising from archesporium. X136; fig. 8, detail of fig. 7, showing me 
mother cell, adjacent tissue dividing, and flattened cells (c) bounding nucellus. 


more vacuolated and which has a large nucleus with the chroma 1 ' 
arranged in a fine network. The cells about it stain rather m<* 
deeply, and toward the chalazal end there are several rows of flatty 
cells (c) which stain still more deeply, and which form a boum *. 
to the ovule in this direction. These rows curve up abov* the '■ 
rogenous group" and meet the epidermis at the point 
integument and nucellus are separated from each other. 


• bete 





In the region of the nucellar apex, the cells are constantly divid- 
ing as the nucellus increases, but this cell activity is not confined to 
that part of the ovule. On either side and below the mother cell there 
is constant division, as is shown in sections through the mother cell 



9, 10. 

"Fig. 9, older ovule, showing megaspore mother cell. X136; fig. 10, 

and in 

J* **<% again 

sections adjacent to it. Often there are three spindles in a 
M thick. This megaspore mother cell enlarges somewhat 

while the nucellus by repeated periclinal 
becoming extended. This changes the broad, 
lectins: portion to a narrower, more pointed 

outline of the 
0nc * The intesra 

development and narrow 






this change in the nucellus the mother cell divides (fig. n) into an 
upper larger and a lower smaller cell. The next stage seen (fig. 12) 
had a row of four megaspores, and the position of the upper two and 
again of the lower two would indicate that they were derived from 
these first two cells by a further division. Fig. 13 shows a disap- 
pearance of the two upper cells, indicated bv the small nuclei and the 

Figs, ii, i 2> 13.— Fig. n, two cells arising from megaspore mother cell; fig- **> 
chain of tour megaspores. X930; fig. 13, four megaspores: three degenerating, the 
fourth the embryo sac. X930. 

narrowed effect of the cells themselves. The nucleus of the cell just 
over the basal one of the chain is being flattened and crushed by the 
development of its sister cell, which is to be the embryo sac & 
other sections there is a deeply stained cap over the micropylar end 
of the embryo sac, which is probably the remnant of the other mega- 
spore or megaspores; at last this also disappears. From this tfo* 
on the embryo sac enlarges rapidly 
(figs. 10, 14). 

and becomes more 





3. Development of the female gametophyte and changes in 

the "spongy tissue" 

Many slides show the uninucleate condition of the embryo sac, 
indicating that this stage is quite prolonged. Fig. 14 shows a few 
cells surrounding the embryo sac, 
which are broken down by the 

growing sac ; material killed August 
29 is about the first to show this 
condition. The nucleus of the 
embryo sac divides as is described 
for other species, and the nuclei take 
the polar position. There is a slight 
indication of cytoplasm forming a 
lining to the wall, but the karyo- 
kinetic figure in the section drawn 
must have formed a little to the 


Figs. 14, 15. — Fig. 14, detail of enlarging embryo sac. Xo3o;"; fig. 15, detail of 
embryo sac and "spongy tissue" (August 29): a, active nutritive cells; b, tissue 
of closely packed cells; c, flattened cells bounding group. X930. 

side of the middle of the sac (fig. 15). 

Fig. 15 shows the sac and 
the surrounding cells on one side as far as the flattened rows (c) 




forming the boundary of the group of cells which we may call the 

"spongy tissue/' using Strasburger's term. These flattened cells 

of the periphery were first seen in the young ovule (fig. 8). The 

"spongy tissue" may perhaps be referred to the mass which originated 

in a single archesporial cell, but the lack of sufficient material prevents 

me from drawing conclusions. 

By the time the embryo sac nucleus has divided, the "spongy 

tissue" shows differentiation. Next to the embryo sac membrane 




Fig. 16. — Polar position of embryo sac nuclei. 


the cells (a) are 
more vacuolated, and often 

into the sac as 
pressure is released on this 
side. Occasionally these 
cells are broken apart and 
the walls are very indis- 
tinct. The next rows (b)- 
often eight or ten in num- 
ber, are made up of smaller 
cells, therefore seeming to 
be closely packed and often 
dividing in such planes that 
they form rows radiating 
from the center of the sac. 
The nuclei and cells them- 
selves stain deeply, con- 

trasting both with the inner layers mentioned and with the flattened 
peripheral rows (c). Quite often a spindle is seen in these cells, so 


that this is by no means a degenerating tissue. The cytoplasm in 
preparations has separated from the walls a little, and the walls ar 
"so transparent that the nuclei seem to be floating in cytoplasm 
(3), but I am convinced that this is due to a slow passage of the 
killing fluid into the tissue and the consequent shrinkage. Fig. 1 
shows the two embryo sac nuclei at the poles, and fig. 17 nas 
nuclei in one section, one of which has not yet passed to the peripi 
eral position. The interior of the sac in these sections has a beau > 
ful vacuolate structure, and the "spongy tissue" has not changt 





nuclei and their peripheral position in the cytoplasmic lining, which 

. h FlGS * *7, 18.— Fig. 17, four nuclei of embryo sac. X930; fig- l8 > embryo sac 
"* layer of cells; spongy tissue beginning to break down. X250. 

•s now a little thicker. The cytoplasm here is quite foamlike in its 
structure (figs. 18, 19). Fig. 19 is a detail of the sac and the "spongy 





The result of the growth of the sac is 


the sac are almost used up (a). Seemingly the wall of each cell is 

--•■« , 

Figs. 19, 20.— Fig. 19, detail of fig. 18: a, "spongy tissue" broken do ** n \^ 
nutritive tissue; 6, compact tissue; c, flattened cells. X930; fig. 20, ovule s 
unusual development of " spongy tissue." 'X37. 

attacked first, probably by an enzyme secreted by the^developi 
gametophyte, and the cells become separated from each ot 
Sometimes a nucleus has its position in an isolated cell, and so 
times it is surrounded by cytoplasm. The breaking down 01 
tissue resembles that described by Chamberlain (4) in the formatio 
of the pollen chamber of Dioon edule. 




Next to this broken tissue there are several layers of cells {a!) which 
resemble those in fig. 16 immediately surrounding the sac. These 
are swollen and have every appearance of being active, nutritive cells. 
Beyond this again is the deeply staining tissue (b) y but narrower now, 
as if some of the cells had changed in their appearance and had taken 

Fig. 21. — Detail of fig. 20. X250. 

on the function of the cells which previously bordered the 
tissue is narrower still toward the chalazal end of the sac. 


made from an ovule in which the embry 


was unusually large and compact. The outer part shows radiation 
n its cell rows, as in other earlier stages. In one case this increase 

great that it almost filled the interior, and 



was so 

the embryo sac had not developed. All these facts point to an active 





one which is formed early and which disappears early, leaving only a 
few thin layers, called "tapetum" by Lang (3) and not explained. 

In fig. 22 this encroachment upon the "spongy tissue" has gone 
on until only about three or four rows are left between the sac and 
the bounding narrow cells (c). These inner cells have enlarged 
many times, as is shown by the magnification of the figures recorded. 
The walls of these cells (b) have become quite thick, the cells are full 










r, ■■ 


1 •© 




* w 




r - 






■ ■ 


Fig. 22. — Detail of embryo sac with many free nuclei; diminished * spong) 
tissue"; cells of tissue (b) have increased in size and assumed nutritive function; 
c, flattened cells. X 580. 

of cytoplasm, and the nuclei are not large, though not disorganized. 
The sac has now a broad cytoplasmic layer and a heavy wall The 


at least 

none were seen in a careful examination with a -fa oil immersion left- 
The membrane is drawn away from the "spongy tissue " for a little 


if this end of the sac was not filled in. 

From this stage on, the history of the development of the sac has 

forms bv Warming 

tion of the endosperm formation is excellent, by Ikeno (7), and b> 
others, so that it did not seem worth while to examine later material, 
at least not until the gaps behind could be closed up. 



1. The young staminate and ovulate strobili can be distinguished 
by the difference in breadth, number of sporophylls, and number of 
meristematic points. 

2. There is probably a single archesporial cell giving rise to a 
group of cells, one of which becomes the megaspore mother cell. 

3. There are four potential megaspores, the lowest one becoming 
the embryo sac, whose development agrees with the accounts of 
other cycads in the main points. 

4. The "spongy tissue" is an active, nutritive tissue, adding to 
its width by division of its cells as it is encroached upon by the embryo 


5. In its final degeneration, the cells of the "spongy tissue" 
nearest the embryo sac are first attacked, and the smaller cells outside 
them take their place, becoming large, swollen, and nutritive in 

Smith College 
Northampton, Mass. 




Smith, Frances Grace, Morphology of the trunk and development of the 

microsporangium of cycads. Box. Gazette 43:187-204. pi. 10. 1907. 

Lang, W. H., Studies in the development and morphology of cycadean 

sporangia. II. The ovule of Stangeria paradoxa. Annals of Botany 14 : 

281-306. pis. 17, 18. 1900. 
3. Treub, M., Recherches sur les Cycadees. III. Embryogenie du Cycas ' 

circinalis. Ann. Jard. Buitenzorg 4:1-11. pis. 1-3. 1884. 
^4. Chamberlaix, C. J., The ovule and female gametophyte of Dioon edule. 

Bot. Gazette 42:321-356. pis. 13-15. 1906. 

5. Ferguson-, Margaret, The spongy tissue of Strasburger. Science N.S. 
18:308-311. I9 o3. 

6. A\ arming, E., Undersogelser og Betragtninger over Cycadeeme. Oversight 
K. Vidensk. Selsk. Forh. pp. 100, 101. 1877. 

7. Ikeno, S., Untersuchungen uber die Entwickelung der Geschlechtorgane 
and der Vorgang der Befruchtung bei Cycas revoluta. Jahrb. Wiss. Bot. 32 : 
5 S 7-602. pis. 8-10. i88q. 



Irving W. Bailey 

The discoloration of sap wood, or "sap stain" as it is commonly 
called, involves the loss or the depreciation in value of large quantities 
of lumber annually. Therefore, with the continually increasing 
value of lumber and the necessity for greater economy in the utiliza- 
tion of wood, the problem of saving and using the sap wood as well 
as the heart wood of the tree increases in importance each year. 
In endeavoring to prevent the discoloration of sap wood it is oi 
fundamental importance to discover what agency or agencies pro- 
duce the stain, and to study their mode of activity. 

So far as the writer has been able to determine, there are in general 
two agencies which produce the discoloration of sap wood. One oi 
these, which is of frequent occurrence and of great economic impor- 
tance, is a purely chemical reaction which takes place in green sap 
lumber upon exposure to the air (oxygen). This chemical discolora- 
tion occurs in many varieties of wood, and is well illustrated by the 
reddish yellow or rusty colored sap stain which occurs in the sap 
wood of alder, birch, and cherry, and by the blue colored sap stain 
which occurs in the sap wood of red gum (Liquid ambar) . When 
the freshly cut surfaces of these woods are exposed to the air, under 
favorable conditions of temperature and moisture, a chemical reaction 
starts which with varying rapidity produces a colored substance i 
the wood. Favorable conditions for sap-staining are found durin 
warm weather, and optimum conditions during extremely ho • 
humid, summer weather, when lumber becomes discolored within 
a few hours. The examination of microscopic sections of this sap 
stained lumber reveals the fact that the colored substance, produce 
by the chemical reaction, is most conspicuously developed in the * 
rays and wood parenchyma cells, living tissues which are largt j 
concerned in the storage and conduction of food in the wood. 


i Contributions from the Phanerogamic Laboratories of Harvard lni\er*i. 

No. 26. 

Botanical Gazette, vol. 50] 



second agency which produces discoloration in sap wood is the activity 
of fungi, which find an abundant food supply in the sap wood, and 
under favorable conditions of heat and moisture (hot, humid weather) 
develop with great rapidity. Thus part at least of the blue colored 
sap stain which occurs so frequently in pine may be produced by the 
dark colored mycelium of a fungus which is found in the food-con- 
taining wood ra^s and the parenchyma cells surrounding the resin 
canals. In fact, in all samples of blue sap pine which have come under 
my observation the color has been produced by this dark colored 
mycelium. However, it seems very likely that a large part of the 
blue sap stain in pine is produced by chemical reactions, for I am 

informed by lumbermen that sap pine lumber frequently stains 
badly during a few hours. The rapidity with which the discoloration 
is produced indicates the activity of a chemical reaction. 

Although the two agencies producing sap stain are so fundamentally 
different, yet the conditions which favor their activity are very similar. 
As has been indicated above, each agency, in its activity, is closely 
related to the food substances contained in the wood. This is shown 
by the fact that the discolorations, produced by the activity of fungi 
and by chemical reaction, are most conspicuously developed in the 
wood rays and wood parenchyma cells. Both agencies producing 
sap stain are, in addition, dependent upon certain quantities of oxygen 
(air), heat, and moisture. Thus optimum conditions for sap-staining 
are found in green sap lumber during hot, humid weather, whereas 
unfavorable conditions are found in cold, dry weather and in logs 
immersed in water. It is well known that certain quantities of food, 
oxygen, moisture, and high temperature are necessary for the rapid 
development of fungi, but it seems advantageous to examine with 
greater care a chemical reaction which is dependent upon similar 

Oxidizing enzymes 

It has been known for many years that certain soluble ferments, 
which facilitate the oxidation of organic compounds, are widely dis- 
tributed in plants and animals. These oxidizing enzymes are of funda- 




produce certain striking postmortem changes in animal and plant 
tissues. I refer in particular to the postmortem discolorations of 
certain organic compounds which are produced by the oxidizing 
activity of these enzymes. The most striking illustration of this 
oxidation and discoloration in plants was pointed out, in 1883, by a 
Japanese, Hikorokuro Yashida, who discovered that the formation 
of black varnish by the oxidation of the latex of the lac tree was 
produced by the intervention of an oxidizing ferment. Since this 
discovery, the activity of oxidizing enzymes has been carefully studied 
in the discolorations which are produced by them in the extracted 
juices of fruits, vegetables, cereals, mushrooms, and other soft plant 
tissues. These investigations have shown conclusively that, as m 
the case of the oxidation of the latex of the lac tree, postmortem 
discolorations are produced in these juices by the intervention or 
oxidizing ferments. This change of color which is produced in the 
extracts of plant tissues has been used as the basis for very delicate 
tests for oxidizing ferments. A strong blue color produced in an 
alcoholic tincture of guaiacum, in the presence of oxygen or hydrogen 
peroxid, indicates the presence of oxidizing enzymes. Upon the basis 
of these tests certain authorities divide oxidizing ferments into 
oxidases and peroxidases, according to whether the blue color 
produced in the tincture of guaiacum occurs in the presence of oxygen 
or hydrogen peroxid. 

It is of interest to note the behavior of oxidizing ferments under 
variations of temperature. Investigation has shown that optimum 
temperatures exist for oxidizing enzymes at which they react wit 
great activity. Below these optimum temperatures their activit; 
decreases, and similarly in increasing the temperature above tne 
optimum their activity decreases, and in almost every case their 
activity is entirely destroyed before a temperature of 100 C. is attainc 
The activity of oxidizing ferments is also decreased or destroje 
by certain antiseptics, and by other chemical substances. 

From this consideration of certain properties of oxidizing enzyme 
we see that there exists a strong similarity between their oxidizm- 
activity and the chemical reactions which produce sap stain, 
each case postmortem oxidization with change of color is produce 
by solutions in contact with the air (oxygen), and variations in tempera- 


ture produce similar variations in the activity of the discoloring 

a 2 


Prevention of sap stain 

The fact that the temperature of boiling water destroys the activity 


sap stain were produced through the action of oxidizing enzymes, a 


be secured by immersing lumber in boiling w r ater. During the spring 
of the present year the writer has tested this method on small boards 
(i"X3"X6") of Alnus incana Moench., Betula populijolia Marsh., 

papyrijera Marsh., and 

These species 

were selected as, at this latitude (Massachusetts), the sap wood stains 
very easily and rapidly during the spring upon exposure to the air, 
whereas the sap wood of such trees as sweet gum, maple, and basswood 
stain deeply during extremely hot and humid summer weather. The 

boards immersed 
permanently unchanged 


whereas untreated pieces stain rapidly and deeply. The fact that 

and that the d 

coloration was prevented by boiling offered strong evidence for believ- 
ing that sap stain results from the action of oxidizing ferments. On 



of oxidizing enzymes. 

In conducting these experiments the boards which were to be 
boiled and those which were to serve as controls were in all cases cut 
from the same part of the same tree, and were subsequently exposed 
to the same atmospheric conditions of temperature and humidity. 
The extreme sensitiveness of oxidizing enzymes in their oxidizing 
activity was clearly demonstrated by the species of wood used in 
these tests. For example, alder and birch, which in hot, humid 
weather stain in a few minutes to a reddish yellow or rusty color, 
stain more slowly at lower temperatures, and in cold dry weather 
stain but slightly even after an interval of several weeks. In a similar 

temperature and moisture. 




whereas boards dried in damp conditions in the open stain very deeply 
and densely- Boards treated in hot water and dried under cover 
remain unchanged in color throughout, and those placed in the most 
unfavorable conditions of abundant moisture and high temperature 
in the open, although they scorch superficially, as does all lumber 
exposed to the direct rays of the sun, remain unstained beneath this 
thin layer of tan. 

Practicability of hot water treatment 

It will occur to some readers that the activity of oxidizing enzymes 
could be prevented by kiln drying or steaming the wood, and that one 
of these methods would be more practicable than immersion in boiling 
water. However, to those familiar with saw mill practice it is evident 
that in mills sawing many thousand board feet of lumber each 
day and night it is usually difficult to steam or kiln dry more than a 
small percentage of the cut. Steaming or kiln drying are expensive 
processes, and the latter can only be applied to certain varieties and 
grades of wood. Furthermore, these processes are time consuming 
and require much handling of the lumber. Some method must be 

devised for preventing sap stain which can be carried out very cheap J 
and rapidly, in order not to interfere with the movement of the lumber 
from the saw to the yard. In other words, the lumber must be 
removed, treated, and piled as rapidly as it is sawed, in order not to 
interfere with the daily output of the mill. Some of the larger mills 
in the south have installed tanks containing chemical substances 
which destroy the action of oxidases and prevent discoloration } 
fungi. The lumber as it comes from the saw is carried through this 
tank upon carrier chains and thus immersed and coated with t e 
chemicals. Certain difficulties have been encountered in the u^ 
of this method. One of the most serious is the fact that the chenuca 5 
used fire proof the outer surface of the boards, and when the treate 
lumber is shipped to northern markets for finishing the planings an 
shavings cannot be burned. This obliges the use of coal to run t e 
planers and other machinery, and the non-combustible planings m 
be disposed of. , 

In the treatment of lumber with boiling water a similar mctno 
could be employed. Long shallow tanks in which water is W* 


by steam pipes or by steam exhausts could be arranged so that the 
lumber from the saw could be passed through the tanks on carrier 

During the coming summer the writer will extend his experiments 
to include a wider variety of woods, and will conduct experiments of 
a practical nature in a large southern saw mill, to determine the relative 
advantages of hot water treatment and treatment with chemical 

Summary and conclusions 

1. Sap stain is in general produced in two ways, by the attacks of 
fungi and by chemical discoloration. 

2. Chemical discoloration is produced in sap wood by the activity 
of oxidizing enzymes. 

3. Hot, humid weather is very favorable to the activity of these 
ferments, and cold winter weather is unfavorable. 

4. Oxidizing enzymes which produce sap stain in wood are 
destroyed and their oxidizing action prevented by a temperature of 



water destroys the oxidizing enzymes in the wood and prevents 

sap stain. 

6. Treating sap lumber in long tanks of boiling water appears 
to be a practical method of preventing sap stain, and to be well 
adapted to saw mill practice. 

In conclusion the writer wishes to express his sincere thanks to 
Professor E. C. Jeffrey for valuable assistance in carrying out this 
investigation, to Professor G. E. Osterhout for advice and sugges- 
tions, and to Mr. W. R. Butler for material of sap-stained lumber. 

Harvard University 




(with one figure) 

The object and knife carrying blocks are long and heavy. These have 
planed surfaces of contact with the running surfaces of the base, instead 
of being mounted on ivory bearings, as is the case in the Thoma instrument. 
When oiled, the blocks arp hpld Kv rsmillai-itv c^ firmlv tn tho tracks that 

the whole instrument can be suspended by either block without breakin 
the contact. In this way rigidity is secured and vibration eliminated, so 
far as the running surfaces are concerned. The friction of running is 
less than in the ordinary Thoma. 

The object clamp is mounted on a solid ball (1.25 inches diameter), 
in a tight socket. This ball can be revolved horizontally through 180 
and vertically through about 45°. It has two orienting levers whose axes 

Botanical Gazette, vol. 50] [M 




of contact with the ball are at right angles to each other, so that definite 
orientation of the object in either of two planes is possible. A third lever 
draws the plate which fits around the " shoulder" of the ball down when 
correct orientation has been attained, and makes the whole absolutely 



The knife carrier revolves horizontally on the column of the knife 
block, and is rapidly adjustable to a height of 1.25 inches by means of the 
screw at the top, the two levers behind clamping it instantly in place. 

Arrangement has also been made for the adjustment of the vertical angle 
of the knife. 

The rigidity and rapidity of the adjustment of this instrument are found 
invaluable in cutting many and long series of wood sections. Its use- 
fulness was demonstrated in the sectioning for Mr. W. P. Thompson's 
work on the rays of the conifers recorded in this issue of the Botanical 
Gazette, and the structure of the instrument is outlined here in the hope 
that others may find it useful. 

No small amount of the credit for the performance of the instrument is 

in the designing. 

workmanship of Mr. H. W. Spexce, y 
R. B. Thomson, University of Toronto. 



The oxidases 

KasTLE 1 has recently published a monograph on oxidases and other oxygen 
catalysts concerned in biological oxidations. The work is a compilation of our 
present knowledge concerning oxidases and other oxygen catalysts, as we as 
an excellent historical resume of the subject. The constantly growing recogni- 
tion of the important role of oxidases and related oxygen catalysts in biologica 
processes, as well as the rapidly growing literature on the subject, makes a sum 
mary of the real contributions especially valuable at the present time. 

The first chapter deals with the important past and present theories of oxida- 
tion, beginning with Schoenbein's ozone theory. After a brief discussion ^ 
this theory, the author proceeds with a more detailed account of Vant HOF 
theory of ionization, Hoppe Seyler's nascent hydrogen theory, and the pero. 
theory of Traube, Engler, and Bach. The conception which involves an 
exchange of electrical potential in oxidations, however, is not mentioned. 

The second chapter takes up the oxidizing ferments, and begins with a 
cussion of their role and range in biochemical processes. This is followed JJ 
detailed account of the guaicum reaction, since our first knowledge of oxi 
ferments is so closely associated with the reaction. The historical trea 
of oxygen exciters and oxygen carriers is divided into tw T o periods. A fte 
covers the first sixty years of the nineteenth century, with Schoexbeins 
tributions standing out as the most important. He supposed that by mea 
various substances and under various influences the oxygen of the air oe 
ozonized. These substances may in turn combine with ozone thus produc 
form an active ozonid, which in turn can give up its oxygen to other less r ' • 
oxidizable substances. Thus the presence of oxygen activators and earners 
recognized, and the most important characteristics of oxidases and peroxi 
discovered, although up to this time these terms had not been introduce 
science. „_ 

During the second historical period, the contributions of Traube an 
trand are"especially emphasized. Traube in his Theorie der Fermentwtrknng ^ 
(1858) established the chemical entity of oxidizing ferments and their ^ 
portance in acting as chemical go-betweens between free or combined 0. . E 

1 Kastle, J. H., The oxidases and other oxygen catalysts concerned in bio ogu^ 
oxidations. U.S. Public Health and Marine Hospital Service, Hygienic Labora o 

Bull. 59. 1909. 



fermentable substances. To Bertrand we owe the introduction into 



the discovery and characteristics of laccase and tyrosinase. The chapter con- 
cludes with a classification of oxidases and special reference to the sources, prepara- 
tion, and characteristics of laccase, tyrosinase, aldehydase, and the purin oxidases. 
The third chapter is devoted to the peroxidases and catalases. The weight 
of opinion is inclined to the conception that peroxidases are substances capable 
of forming unstable peroxids from hydrogen peroxid, by double decomposition 




Bach and Chodat's conception of an oxidase consisting of a mixture of 

peroxidases and peroxid-forming substances (oxygenases) would make the 

peroxidases the more important agents in plant and animal oxidations, and would 

relegate the oxidases to an insignificant position in such oxidations, if indeed 

they function as enzymes at all. In the author's opinion the objections which 

have been recently urged against the true enzymatic nature of oxidases are well 

A considerable amount of evidence is brought together to show the importance 
of iron, copper, and manganese as coenzymes to oxidizing ferments. According 
to Bertrand, manganese is the really active element of the oxidases, so far as 
the activation and transfer of oxygen is concerned. Euler and Bolix have 
found that laccase has no action on hydroquinone in the absence of manganous 
salts, and therefore they suggest that laccase owes its activity to the presence of 
such salts. In this connection it is interesting to note that in a paper by Bach, 2 
more recent than the above monograph, he claims to have obtained a tyrosinase 
which will oxidize tyrosin to the red stage and is free from both iron and man- 
ganese. He concludes, therefore, that manganese and iron salts are in no way 
necessary for oxidase activity. One of the most valuable features of the mono- 
graph is the comprehensive list of references to the literature.— Charles O. 

Colloidal chemistry 

The newer plant physiology should welcome the appearance of Freuxdlich's 
book 3 on colloidal chemistry, or capillary chemistry, as he terms it. This is the 
first attempt to bring together our knowledge of this youngest and most difficult 
branch of physical chemistry. It puts the physiologist immediately in touch 
with the present status and most important literature of a subject which seems 
destined to play at least as important a role in the study of vital phenomena as 

2 Bach, A„ Zur Theorie der Oxidasewirkung. Ber. Deutsch. Chem. Gesell. 
43:362. 1910. 

* Freuxdlich, Herbert, Kapiiiarchemie, eine Darstellung der Chemie der 
Kolloideund verwandter Gebiete. 8vo. pp. viii + 591. figs. 75- Leipzig- X9°Q- 


has the chemistry of true solutions. The importance of surface tension in the 
mechanics of cell life has been emphasized, adsorption phenomena have recently 
become important in our consideration of semipermeability, selective absorption, 
etc., they must also be called upon to explain many of the relations between soil 
and plant. The student of protoplasmic structures, nuclear membranes, chromo- 
somes, fibrillae, and the rest, is perhaps more familiar with coagulation of colloids 
and adsorption than he knows; his killing and fixing are examples of the former, 
while the whole process of staining apparently depends upon the different adsorp- 
tive powers exhibited by various portions of the coagulated protoplasmic mass. 
The so-called Brownian movement and the other phenomena of motion usually 
observed in the protoplasmic emulsion are likewise to be classed under capillar} 
chemistry. If enzyme action is to receive an explanation, it bids fair to come 
also from this realm. 

With this book and the field that it represents once in general use, it would 
seem that physiological research should receive a very great impetus along just 
those lines where it now wavers most. One of imagination, who appreciates the 
problems and present rapid advance of this and other branches of physical chem- 
istry, should have little cause so thoroughly to lose heart as to need the aid ot 
those "entelechies" and other dei ex machina with which the "neo-vitalism 
seems to be somewhat overburdening biological philosophy. In the present 
exposition of capillary chemistry the author proceeds from the simpler phenomena 
of surface tension and capillarity to subjects of more complex nature, such as 
adsorption, colloidal solutions, suspensions, emulsions, catalysis, and the like. 
Every section is brief, clear, and directly to the point in hand; experimenta 
evidence is given prominence rather than theoretical deductions, though the 
latter are not wanting; and numerous footnotes orient the reader in the scatters 
literature of the subject. An index of authors and one of subjects enhances the 

value of the work. — B. E. Livingston. 


Insect galls .—Miss Stebbins 4 has published a bulletin on insect galls of 
Springfield, Mass., and vicinity, which will be very useful to botanists who are 
interested in cecidology. The galls are grouped with reference to the plants, 
which have been arranged in accordance with Brixton's Manual. This is t e 
first American work in which these pathological growths have been groupe 
with reference to the host plants. The record shows 204 species of gall -producing 
insects, which are distributed in 52 genera, 14 families, and 6 orders. The ga s 
occur on 93 species of host plants, which are distributed among 48 genera, 29 
families, and 16 orders. The descriptions of the galls are clear and are reinforc 
by 1 12 illustrations. The descriptions of the insects are omitted, but the synonom) 
and bibliography given with each will enable the student of entomology to loo- 
them up without difficulty. The galls of 26 new species are described and nam , 

4 Stebbins, FANNER A., Springfield Museum of Natural History, Bulletin 2. 1Q 10 ' 


and 6 are described without names. The describing and naming of new species 
of galls without the insects has been the subject of considerable criticism, but 
since it gives us a definite record of these species the reviewer is inclined to favor 
the violation of this law of nomenclature. The work closes with an extensive 
bibliography, a systematic index of gall insects, and an index of scientific and 
common names of host plants. — Mel T. Cook. 



Influence of environment on wheat. — One of the most persistent theories in 
evolutionary discussions of cultivated plants is that of the " breaking up of types, 
supposed to be brought about when plants are grown from seed under conditions 
differing markedly from those under which the parent plants were grown. Evi- 
dence for this view has been largely of an observational nature and capable of 
other interpretation. Experimental evidence bearing on the question has been 
brought out by LeClerc and Leavitt 5 in reporting a series of cultures of 
wheat in widely different sections of the United States. The plan of the experi- 
ments was as follows: In one series Kubanka wheat grown in South Dakota was 
distributed to stations in Kansas and California, a sample being likewise grown 
in South Dakota. Every year a sample from each station was sent to each of 
the others and grown there. A similar series of cultures was 
Crimean wheat in Kansas, Texas, and California. Some of the experiments 
have now been continued for five years. 

The results may be briefly summarized. The original pure type of Kubanka 
wheat from South Dakota showed entirely different morphological characteristics 
and chemical composition at the different stations. The characteristics of the 

carried out wi 



derived. Thus, when South Dakota wheat was grown in Kansas or California, 


peculiar for each region; but if, after several generations, these wheats were 
again transferred to South Dakota the resulting crop assumed all the characteristics 
of the same variety grown continuously in South Dakota. The series with 
Crimean wheat gave exactly similar results. 

The experiments show that wheats of one variety from several sources, when 
grown in the same locality, differ but little in morphological characteristics and 
chemical composition, but if grown in different localities from seed of the same 
source, they differ widely from each other. There is a marked response to 
environment, but all the plants of a pure variety respond in the same way. There 
is no tendency toward "breaking up" of the type on account of change in environ- 
ment— H. Hasselbring. 

5 LeClerc, J. A., and Leavitt, S., Tri-locai experiments on the influence of 
environment on the composition of wheat. U.S. Dept. Agr., Bur. Chem., Bull. 128. 
PP- 18. i 9io . 


Alcoholic fermentation. — Kohl 6 has carried on a study of the series of 
reactions involved in alcoholic fermentation. He finds that lactic acid is not 
fermented either by zymase, compressed yeast, or bottom yeast; that i per cent 
or more of lactic acid stops the self-fermentation of living yeast and strongly 
reduces its fermentative activity in glucose; but that zymase, compressed yeast, 
and brewer's yeast ferment sodium lactate speedily. It is evident that if at one 
stage of alcoholic fermentation lactic acid is found, it must exist as a salt. The 
fact that zymase will not ferment lactic acid has been urged against Buchner s 
conception that alcoholic fermentation takes place in two steps; glucose is trans- 
formed to lactic acid by zymase, and lactic acid to alcohol and carbon dioxid by 
lacticidase. Kohl's finding answers this argument. While he thinks the fer- 
mentation occurs in these two steps, he differs in his view of the enzymes that 
carry on the processes. He concludes that catalase transforms the glucose to 
lactic acid, and that zymase carries the splitting on to alcohol and carbon dioxid. 
In a glycerin extract of crushed yeast, he found neither an oxidase nor a peroxidase, 
as shown by an alcoholic solution of quaiac; yet it contained an enzyme capable 
of oxidizing various phenols, and these oxidations he believes are carried on bj 
the catalase present. This extract when filtered produces lactic acid in the 
presence of glucose — it likewise produces a trace of oxalic acid. He does no 
know whether both these oxidations are due to the same enzyme. He urges 
that this view locates the function of the catalase of yeast, a point not before 
settled. In case zymase is present, the oxidations go no farther than lactic aci , 
which is then transformed to alcohol and carbon dioxid. In its absence, the 
oxidation is carried still farther, producing various other acids. He apples his 
view to the explanation of the results of Harden and Young with the gelatin 
filter, but cannot be said in any degree to further elucidate them. This subjec 
because of its close bearing on respiration and energy-production in the organis 
certainly needs much attention from biological chemists. It is disappointing, 
however, that the contributions are mainly hypotheses with sparse experimen 
evidence, rather than records of careful chemical studies.— William Crocke 


Adoxa moschatellina. 

positio- -' AdMM induced 

Lagerberg 7 to undertake a complete morphological and cytological investiga 
tion of this peculiar genus. The development of the various organs was trace * 



are illustrated by three large plates. The following are some of the princip* 
features: The ovule has a single integument and a single archesporial cell v» 


6 Kohl. F. G m Ueber das Wesen der Alkoholgarung. Beih. Bot. Centralb . 

2Q i :\ 15-126. 1910. 

7 Lagerberg, TV, Studie 

Stellung von Adoxa moschatellina. Kungl 
figs. 23. 1909. 

ien iiber die Entwickelungsgeschichte und systema 
ellina. Kungl. Sv. Vet. Akad. Handl. 44 -^ 86 - &*' 


develops directly into the embryo sac, as in Lilium, the four megaspores, not 
separated by walls, all taking part in the formation of the sac. The two male 
cells retain their form even after passing to the end of the pollen tube. Double 
fertilization was observed. The diploid and haploid chromosome numbers are 
38 and 18. The first four cells of the endosperm are long and tubelike, extending 
from the egg to the antipodals. 

Various genera of the Saxifragaceae, Araliaceae, Caprifoliaceae, and Ranun- 
culaceae, with which various systematists have supposed Adoxa to be related, 
were studied for comparison, and one of these genera (Sambucus) shows so many 
resemblances that the similarity could hardly be accidental. For instance, both 
have ovules with a single integument and a single archesporial cell which develops 
into the embryo sac according to the Lilium type; the wall of the anther, the 
cytological details of the development of the pollen (including the number of 
chromosomes), and the structure of the mature pollen grain are so identical that 
the two forms can hardly be distinguished in these respects; the long persistence 
of the organized male cells is the same in both, and resemblances in the grosser 
morphology were already well known. The conclusion is reached, and it seems 
to be based upon an unusually wide range of evidence, that there is no need 
for the family Adoxaceae, and that Adoxa should be placed in the Caprifoliaceae 
in the tribe Sambuceae — Charles J. Chamberlain. 

Geotropism.— Under a very pretentious title, Giltay 8 discusses and 
describes a number of experiments on some of the "fundamental questions of 
geotropism." The article is more a contribution to the teaching of the subject 
than to knowledge. He gives an excellent method for lecture demonstration 
of the force with which a geotropically bending primary root turns downward. 
He also points out that we have never proved that gravity is the only stimulus 
involved in the turning of the primary root toward the center of the earth. The 
only evidence we have for this is qualitative. Knight showed (1806) that as 
the centrifugal force was increased on a centrifuge with a vertical axis the root 
and stem took more nearly the horizontal position; but he did not show any 
relation between the position of the stem and root and the resultant of the two 
forces which he assumed to be involved (gravity and centrifugal force). In 
short, Knight showed that at least in part the so-called geotropic stimulus is 
the gravity stimulus, but he did not show that the gravity stimulus is the only 
stimulus involved. Giltay urges the necessity of showing that the position 
taken by orthotropic organs on such a centrifuge is the resultant of the two forces, 
,f we are to be assured that the geotropic stimulus is identical in nature with 
the gravity stimulus and with the stimulus of centrifugal force. The resultant 


..w tunc* uugni 01 course to be tne position taKen, proviucu u^ u»v, w*v,~ 
(| o not differ from each other by many fold, for we must remember that Weber's 

8 Giltay, E., Einige Betrachtungen und Versuche iiber Grundfragen beim 
eotropismus der Wurzei. Zeitschr. Botanik 2 :30 5 -3 3 i. 1910. 


law applies in geotropism as in various tropic responses. Giltay devised a 


special centrifuge with a vertical axis for testing this point. He 
the average of 368 tests, with the angle of the resultant approximating 45°> the 
primary root fell 2 . i° below the resultant. This deviation can well be accounted 
for by the variation in speed of rotation and the variation of roots themselves. 
This seems to furnish evidence for the identity in nature of the geotropic and 

centrifugal stimuli. — William Crocker. 

Role of hydrocyanic acid— Treub^ has found that the amount of hydro- 
cyanic acid in plants of Sorghum increases during the day, not due to the direct 
action of light, but in proportion to the formation of the products of the assimila- 
tion of carbon. It was already known, from investigations with Pangium edule 
and Phaseolus lunatus, that light has no part in the formation of hydrocyanic 
acid except as it favors photosynthesis. Much the same results have been obtained 
with Primus javanica. Passi flora joetida and at least four other plants offer 
examples for the demonstration of the direct proportion between the formation 
of hydrocyanic acid and the function of chlorophyll. This can be demonstrated 
also by the use of variegated leaves. The amount of acid is usually greatest 
in the young leaves and gradually diminishes as the leaves grow older. l 
Sorghum, young leaves grown in a dry season or on dry soil contain much ac , 
and for this reason are dangerous as food for stock. Leaves about to fall con 
very little acid, while, with only two exceptions, those already fallen contain no . 
Guignard found that fallen leaves of Sambucus nigra contain much of the aci . 
Treub confirms these results and finds the same to be true of fallen leaves 
Indigofera galegoides. The hydrocyanic acid is probably the first recogniza 
simple organic product of the assimilation of nitrogen, and perhaps the 
organic nitrogen compound formed. The amounts of the acid in plants wate 
with a solution of sodium and potassium nitrate increased or decreased in p 
portion to the amount of nitrate used. Ravenna and Peli think that nitra 
and carbohydrates are necessary to the formation of the acid. Treub agre 
with these conclusions, and adds that dextrose is especially essential- 
acid probably does not occur in plants as such, but in the form of a glucoside 
which it can be liberated by an enzyme or by boiling water. — R. Catlin K 

Parasitic flagellates in plants. — Although rapid progress in the sfu J 
parasitic flagellates has shown them to be of widespread occurrence in am 
organisms, the discovery of these parasites in plants is a noteworthy fact, 
occurrence of a trypanosome-like parasite in the latex of Euphorbia piluh/er 
Mauritius was first reported by Lapont. 10 The discovery was soon afters 


Treub, M., Nouvelles recherches sur Ie role de I'acide cyanhydnque 
les plantes vertes. Ann. Jard. Bot. Buitenzorg II. 8:84-118. 191 o. 

1° Lafont, A., Sur la presence d'un parasite de la classe des FlageUes dans 
latex de I 1 Euphorbia pilulijera. Compt. Rend. Soc. Biol. 66:1011-1013- J 9°9- 


confirmed by Donovan 11 by observations on the same species of Euphorbia in 
Madras. Lafont 12 now follows with a full account of the organism. The 
parasite, which was originally discovered in the latex of Euphorbia pilulifera, 
occurs also in the two other species E. thymijolia and £. hyperici folia. A search 
of the latex of some 50 other species of plants from various families failed to 
reveal similar organisms. About one-third of the Euphorbia plants from different 
stations were found to be infected. The number of parasites in different plants 
varies greatly. The infected plants show the effects of malnutrition, and finally 
drop their leaves and die. The protozoans are elongated, flattened, and some- 
what undulate. They do not, however, possess the undulating membrane of 
trypanosomes, and are therefore placed in the genus Leptomonas, as L. Davidi. 
The apex is provided with one cilium, which originates in a blepharoplast. 
A large nucleus is situated near the center of the body. Division, which was 
observed in hanging drop cultures, takes place by longitudinal fission, preceded 
by a thickening of the body of the organism. Various forms, perhaps indicating 
different stages in the development of the organism, were observed. The sim- 
plest are spherical, nucleated masses of protoplasm, which soon form a cilium. 
It is possible that two parasites exist here. Injection of the parasites into the 
blood of small animals produced no infection, although some of the animals 
died from unknown causes.— H. Hasselbring. 

Diseases of celery. 


tributions of life histories of Fungi imperjedi an account of two diseases of celery 
occurring in the truck gardens on the lowlands surrounding Hamburg. The 
first is the leaf-spot disease caused by Septoria Apii (Briosi and Cav.) Rostr., 
also known as 5. Petroselini Desm. var. Apii, and as Phlyctaena Magmtsiana 
(Allechr.) Bres. The fungus attacks the leaves, stems, and fruits of the celery 

In following out the manner 


m which the fungus lives through the winter, the author encountered no other 

Minting stages. The fungus is carried over from year to year by means ot spores 
which persist both in the pycnidia on the plant remnants left in the fields, and 
m the pycnidia on the seeds. With spores from both sources the author was 
able to produce infections on young plants with ease. 

The second disease is a scab of the roots, which, although it has been reported 
from several places, has never been critically studied. The disease is shown to 



11 Donovan, C, Kala-azar in Madras, especially with regard to its connection 
With the dog and the bug (Conarrhinus). Lancet I77:i495-i49 6 - i9°9- 

12 Lafont, A., Sur le presence d'un Leptomonas, parasite de la classe des Flagelles 
1 ans le latex de trois Euphorbiacees. Ann. Inst. Pasteur 24:205-219. figs. 7. 1910. 

^Klebahn, H., Die Krankheiten des Selleries. Zeitschr. Pflanzenkrank. 20: 
^40. pis. 2. figs. 74 lgi0 . 


The pycnidia occur on the diseased roots, and more abundantly on the lower 
part of the petioles and on the fruit, but rarely on the leaves. Cultures were 
obtained from hyphae invading the sound tissue of the roots and from spores. 
The colonies from both sources were similar, and many infection experiments 
with mycelium from both sources on sound roots were successful. The action 
of this species of Phonia in producing a scab and rotting of celery tubers is a 
case analogous to the well-known root rot of sugar beets caused by another 
species of the same genus, Ph. Betae.—H. Hasselbring. 

Treatment for smuts.— The usual methods of treating seed-grain for the 
prevention of smuts have not proved applicable in the case of the loose smuts 
of wheat and barley, since these fungi persist through the winter, not by 
means of spores adhering to the surface of the grain, but by means of a dormant 
mycelium in the interior of the seed. Appel, T * following out the suggestion made 
by Jensen at the time of the publication of his hot water treatment to use a 
preliminary treatment with cold water for seed infected with these smuts, has 





at temperatures which do not injure the seed. The experiments substantiate 
this belief. It is found that grain infected with Ustilago tritici or U. nuda can 
be treated successfully by being soaked for six hours at 20-30 C, and by being 
treated subsequently with hot water at 50-54 C, or by hot air at a corresponds ~ 
temperature.— H. Hasselbring. 

Dehiscence of anthers.— Hannig *5 takes up what apparently he regards 
as a real difference between Steinbrinck's cohesion theory and Schneider's 
Schrumpjangstheorie for the explanation of the dehiscence of anthers. To the 
reviewer the two theories differ more in the degree of analysis than anything else, 
as he believes that this phenomenon must be in the last analysis found dependent 

the tensile strength of water. However, the author has done a real service 
in showing how the dehiscence is a genuine cohesion consequence. He has 
accomplished this by artificially causing dehiscence through the effect of dehy- 
drating solutions. He has shown that dehiscence will occur in a saturated atmos- 
phere if anthers are exposed to light which generates enough heat in the tissues 
to reduce the vapor tension sufficiently to set up tension in the water contained in 
the membranes. Burck's notion that the nectaries withdraw water from the 
membranes and hence cause dehiscence in a saturated atmosphere was not con 
firmed. — Raymond H. Pond. 


14 Appel, Otto, Theorie und Praxis der Bekampfung von Ustilago tritici und 
Ustilago nuda. Ber. Deutsch. Bot. Gesell. 27:606-610. 1910. 

•5 HANNIG, E., Ueber den Offnungsmechanismus der Antheren. Jahrb. ^ I5S - 
Bot. 47:186-218. 1909. 


Plant diseases. — Whetzel and Stewart, 16 contrary to the common belief, 
advocate the cultivation of pear orchards if a crop of fruit is desired. In this 
view they are upheld by Hedrick of the State Station, who found that blight 
epidemics are not necessarily dependent upon cultivation and manuring. No 
immunity to the disease was obtained by the use of certain blight remedies. 

Sackett 1 ? has described the appearance of this new bacterial disease in the 
field, and has given the manner of infection, together with a complete morphologi- 
cal, cultural, physical, and biochemical description of the causal organism, 
Pseudomonas medicaginis sp. n. The work is well supported by numerous 
inoculations. The only thing lacking in this well-balanced investigation is a 
bibliography. — Venus W. Pool. 

Source of nitrogen for molds. — Ritter 18 finds that the ammonium salts of 
mineral acids as the source of nitrogen for the molds is inverse to the strength of 
the acid forming the negative ion of the salt. The author attributes this to the 
toxic effect of the acid liberated by the assimilation of the ammonium ion. For 
instance, mono-ammonium or diammonium phosphate is a far better source ot 
nitrogen than the ammonium salts of sulfuric, hydrochloric, or nitric acids. The 
so-called "Nitratpilze" (Aspergillus glaucus, Mucor racemosus, Cladosporium her- 
barium) gave on the average a greater yield of organic material from the two 
ammonium phosphates mentioned than from potassium nitrate. The yield from 
the ammonium salts of the stronger mineral acids was very much lower.— Wil- 
liam Crocker. 

Excretion of salts by Statice — Schtscherback 10 has investigated the excre- 
tion of salts by the leaves of Statice Gmelini. Many leaves of halophytes are 
known to excrete salts in considerable quantities by means of the glands described 
by DeBary and others. Leaves of Statice Gmelini, floating on pure water, are 
soon freed from their contained salt and thereafter excrete water only. The amount 
of excretion of a leaf floating on a solution of a substance depends upon the sub- 
stance and the concentration of it used; sulfates and chlorids of sodium, potassium, 
and magnesium tending to increase it, while calcium compounds and sugars de- 
crease it. The amount of excretion does not depend upon the turgor pressure 
J n the leaf cells.— R. Catlix Rose. 

Physcia villosa in North America.— In a recent number of this journal (49: 
320. i 9I0 ) I recorded this plant from southern California. Since then I have 

16 Whetzel, H. H., and Stewart, V. B., Fire blight of pears, apples, quinces, 
et c Bull. \\Y. Cornell Exp. Sta. 272:31-51. figs. 5-231. 1910. 

»> Sackett, W. G., A bacterial disease of alfalfa. Bull. Colo. Exp. Sta. 158:1-32, 
P ls - *-J. 19 10. 

'* Ritter, G., Ammoniak und Nitrate als Stickstoffquelle fur Schimmelpilze. 
er ' DeutS(:h - Bot. Gesell. 27:582-588. 1909. 


«<«< Gmelini. Beih. Deutsch. Bot. Gesell. 28:30-34. 1910 



found in the herbarium of Wellesley College a specimen distributed with Evernia 
jurfacea (L.) Mann, collected by Edward Palmer at San Diego, California, 
in December 1888. A duplicate of this collecting has been kindly sent me by 
Dr. L. W. Riddle, who also calls my attention to the fact that this plant was 
distributed in Decades North American Lichens (no. 154) from San Quintin 
Bay, Lower California, Mexico, where it was collected by C. R. Orcutt (see 
Hasse, Bryologist 13:61. 1910). — R. Heber Howe, Jr., Thoreau Museum, 
Cancord, Mass. 

Fertilization in Rafflesia.— The remarkable and renowned Rafflesia has long 
attracted attention, but little has been known of its more minute details. An 
investigation 20 of its embryo sac and fertilization shows that in spite of the para- 
sitic habit and grotesque appearance, the development of the embryo sac and the 
process of fertilization are quite normal. It was noted that young stages in the 
development of the ovule are found in nearly mature buds, and that the develop- 
ment of the sac takes place after the flower is open. — Charles J. Chamberlain. 

Microchemistry of chromosomes. 21 — The title arouses interest, but from the 
paper we learn only that chromosomes may be dissolved in hot water, while the 


reticulum of the resting nucleus is little affected, and that therefore the importance 
of chromatin in heredity has been overestimated. That there are chemical changes 
as chromosomes are developed from a reticulum has been known for some time, 
but we now know the effect of hot water upon chromosomes and theories of hered- 
ity. — Charles J. Chamberlain. 

Absorption of salts by Bromeliaceae — From his work with the Bromeliaceae, 
Aso 22 concludes that Ananas sativus, Pitcairnia imbricata, and Nididaria pur- 
purea do not take up, or only in very small amounts, by means of the scales 
the leaves, salts soluble in water. On the other hand, Tillandsia usneoides, 
after five days of submergence in a 0.3 per cent lithium nitrate solution, showe 
in different parts of the plant considerable quantities of the salt— R. Catlin 


20 Ernst, A., und Schmid, Ed., Embryosack entwickelung bei Rafflesia Raima B . 
Ber. Deutsch. Bot. Gesell. 27:176-186. pi. 8. 1909. 

2 * Nemec, B., Zur Mikrochemie der Chromosomen. Ber. Deutsch. Bot. Gesell. 
2 7:43-47- 1909. 

« Aso, K., Konnen Bromeliaceen durch die Schuppen der Blatter Salze au - 
nehmen? Flora 100:447-449. 1910. 

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Botanical Gazette 




Oswald Schreiner and J. J. Skinner 

(with ELEVEN figures) 

The investigation of infertile soils from various parts of the 
United States has received considerable attention in the last few 
years, and has been conducted along several converging lines in 
these laboratories. Among these is a thorough inquiry into the 
nature of the organic matter of soils. The results of these researches 
into the chemistry of the organic matter of the soil, its origin, 
transformation, and properties, have been reported upon to a 
large extent in a former bulletin of this Bureau and in scientific 
journals. 2 Several bodies have been isolated from such soils, which 
have quite different chemical properties, thus showing that there 

1 Published by permission of the Secretary of Agriculture. 

2 Schreiner, O., and Shorey, E. C, The isolation of dihydroxystearic acid from 
soils. Jour. Amer. Chem. Soc. 30:1599. 1908. 

, The isolation of picoline carboxylic acid from soils and its relation to soil 
fertility. Jour. Amer. Chem. Soc. 30:1295. 1908. 

-, The presence of a cholesterol substance in soils; agrosterol. Jour. 
Amer. Chem. Soc. 31:116. 1909. 

, A wax acid from soils; agroceric acid. Science N.S. 28:190. 1908., 

, Pentosans in soil. Science N.S. 31 :3o8. 1910. 

, Purine bases in soils. Science N.S. 31:309. 1910. 

, The presence of secondary decomposition products of proteids in soils. 
Proc. Amer. Soc. Biol. Chem. 1:47. 1907. 

See also Bull. 53, Bureau of Soils, U.S. Dept. Agric. 

Schreiner, O., and Sullivan, M. X., Soil fatigue caused by organic compounds. 
Jour. Biol. Chem. 6:39. 1909. 

• 161 


is a difference in the nature of the chemical bodies in different 
soils. Some of the bodies isolated contained nitrogen, while others 
free from this element contained carbon, hydrogen, and oxygen 
only. The presence of sulfur and phosphorus compounds has also 
been strongly indicated by these studies, although the actual 
identification of bodies of such nature has not been definitely accom- 
plished. The results are sufficient, however, to show that a wide 
range of bodies of different composition exists in soils, and a more 
detailed knowledge of their nature and properties seems imperative 
in order to know and understand the nature of the material influ- 

encing crops in 

microbiologic and enzymotic 

In an earlier paper 3 it was demonstrated that the roots of plants 
possessed a very appreciable oxidizing power. This oxidizing power 
of the normal root was found to be influenced by the medium in 
which it grew. Thus good soils and their aqueous extracts pro- 



the roots. Some substances harmful to plant growth were 
found to have an inhibitive effect on this oxidation. When smaller 
amounts were present, however, it was found that the oxidizing 


An examination in one case, namely that of vanillin, where i 
was possible to demonstrate by colorimetric test the presence 
or absence of minute amounts, showed that this disappear 
entirely from solution under the influence of the oxidizing power o 
the roots. It was further shown that the fertilizer salts, in addition 
to promoting plant growth, also had a very strong influence in pro- 
moting this enzymotic effect, this being especially marked in the 
case of sodium nitrate and lime. 4 The action of fertilizer salts 
and the influence of such harmful soil constituents were further 


Schreiner, O., and Reed, H. S., Studies on the oxidizing power of roots. • 

_rrE 47:355- 1909. See also Bull. 56, Bureau of Soils, U. S. Dept. Agnc. i^9- 

4 Schreiner, O., and Reed, H. S., The power of sodium nitrate and ca | c ^ g 

to decrease toxicity in coniunction with nlants crowing in solution cu 

carbonate rf §m 

Jour. Amer. Chem. Soc. 30:85. 1908. 


other factors reported in this paper. All of the harmful bodies 
which had been studied and presented in previous papers had a 
distinct influence on the roots, causing them in many cases to be- 
come stunted or swollen and darkened at the tips, or to show other 
physiological irregularities of the same kind as is exhibited by the 
roots in different extracts from infertile soils. It is, therefore, of 
interest to know further the influence which such altered root 
conditions would have upon the composition of the soil solution 
and the influence of added fertilizers. 

One of the bodies isolated from a number of unproductive soils 
is a definite crystalline body identified as dihydroxystearic acid 
melting at 98°-99° C. It can be prepared by the oxidation of 
elaidic acid in the laboratory. In these experiments the dihy- 
droxystearic acid used had been prepared in this manner. The 
frequent occurrence of this body in such soils and its disappearance 
therefrom by processes which promote aeration or oxidation made 
this especially suitable for further study of its effects on plant 
development in relation to the concentrations of soil solutions or 
fertilizer application. 

The dihydroxystearic acid can be isolated from a soil containing 

it by treatment with a 2 per cent sodium hydroxid solution, and, 

after allowing the mineral material to settle, the alkaline extract is 

separated and made acid with a slight excess of acetic acid. The 

so-called humus precipitate which is thus formed is filtered off and 

the clear filtrate is shaken out with ether and the ether solution 

allowed to evaporate on the surface of a small quantity of water. 

The dihydroxystearic acid is left on the surface of the water, 

together with other impurities extracted by the ether. The 

impurities can be largely removed by heating the water to boiling, 

and filtering. Fig. x shows the effect of a solution of this nature 

on wheat seedlings when the material is dissolved in much water. 

The dihydroxystearic acid, when dissolved in a small volume of 

water and then cooled, crystallizes out in the form of small plates 

or leaflets arranged in radiating clusters. Fig. 2 shows the effect 

of the purified substance in various concentrations. The details 

of the method of isolating and purifying will be found in the paper 




Dihydroxystearic acid can also be prepared in the laboratory 
by starting with oleic acid, which by treatment with nitrous oxid 
is changed to the isomeric elaidic acid. The elaidic acid thus 
formed is dissolved in a solution of potassium hydroxid and 
oxidized by a solution of potassium permanganate, one of the 
products under suitable conditions being dihydroxystearic acid. 




\ b 







i Hi 





Vi J f*j 


Fig. i.— Wheat seedlings grown in extract obtained in the method for isolating 
dihydroxystearic acid from soils: j, 2, undiluted extract; 3, 4, one part of extract, 
one part of distilled water; 5, 6, one part of extract, nine parts of distilled water. 

Care has to be taken that the oxidation does not proceed too far, 



is very 

readily oxidized to other compounds. 



soil constituent upon plant growth and upon soil solutions i 
fertilizer action, especially with reference to the ratio of phosphate 
nitrate, and potash originally present and removed by wheat seed- 
lings in the course of the experiment. 



Solution cultures 


the three fertilizer ingredients, 

namely P 2 5 , NH 3 , and K 2 0, as calcium acid phosphate, sodium 
nitrate, and potassium sulfate, respectively, in all possible ratios 
of one, two, and three constituents, varying them in stages of 10 
per cent, were prepared, the concentration being 80 parts per 
million in these constituents. In a similar set of cultures there 



Fig. 2. — Wheat seedlings grown in solutions of dihydroxystearic acid from soils: 
1, solution of dihydroxystearic acid, 200 parts per million; 2, 100 parts; 3, 50 parts; 
4, 20 parts; 5, control in distilled water. 


culture. The selection of the salts as carriers of the phosphate, 

terms P„0„ NH 

with the practice in fertilizer 


be seen, are also carriers of calcium, of sodium, and of sulfate, and 


selected for giving at the 
The details of this method of 

same time 

perimentation have been given in 


an earlier paper, and the reader is referred to this for a full explana- 
tion of the use of the triangular diagram and the results obtained by 
growing seedling wheat in these various culture solutions without the 
presence of any added harmful substance. s Wheat seedlings were 
then grown in these various cultures and observations made in regard 
to general development, the effect on the root growth and appearance, 
and on root oxidation, and at the termination of the experiment the 
green weight of the plants was taken. The solutions were changed 



and potash being determined, thus giving the concentration of 
these elements and their ratios existing at the end of every three- 
day period for comparison with the original concentration and 
ratio. This changing of the solutions was kept up for twenty- 
four days, thus making eight changes. 

In the present work a triangle of solution cultures similar to 
the one described in the previous paper was set up, with the differ- 
ence that 50 parts per million of dihydroxystearic acid were present 
in each culture. This set of cultures grew from April 2 to April 20. 
The set without the dihydroxystearic acid grew from February 20 

to March 21. While 


v^****^ ww W..&W iu\.t wnut i*u.V'y n^/iv wivnu Ut umviw^v ~ / 

under very similar greenhouse conditions and for the same lengt 
of time as well as in the same time of year, one closely following 
the other, the results show, nevertheless, very strikingly the effec 
of dihydroxystearic acid, and are duplicated or substantiated by 
two further experiments in which the sets were grown simultane- 

The analytical results, however, were not so complete in these 
later tests, owing to the inability to handle the 396 separate deter- 
minations necessary every three days. The results given in tht> 

^ , , , are largely based on the first set, for the reason 

that the analytical results were more complete than in the other 





stearic acid and those without this substance was very marke ■ 

5 Schreiner, O., and Skinner, J. J., Ratio of phosphate, nitrate, and potassium 
on absorption and growth. Bot. Gazette 50:1. 1910. 


and was especially striking in those sets where both triangles of 
cultures were grown simultaneously. In addition to the general 
appearance of the tops, the presence of this harmful body produces 
still other effects readily recognized by the -investigator, and form- 
ing on the w r hole a better physiological indicator of the toxicity 
than the growth of tops, as has been well recognized by physi- 
ologists generally in conducting similar work. Reference is here 
made to the action of the body on root condition and growth. 
The root being bathed by the solution, and being, moreover, the 
delicate mechanism of absorption, is often a more sensitive indi- 
cator than is the growth of the top. The most marked effects of 
dihydroxystearic acid on roots are strongly to inhibit their growth 
and to produce enlarged or swollen tips, which are frequently very 
dark in color and often turned back in the form of hooks. These 
phenomena are observed even when the injury is not so great as to 
kill the plants. This action of the body on the roots is influenced 
by the conditions of growth imposed upon the plant by the different 
fertilizer ratios. This is shown by the general notes taken on one 
of these sets one week after growth in the cultures had begun. 


.0 nitro 
made 1 


number turned upward. The tops have made some growth, some 



to the above; the roots and tops are somewhat better, but still 
show an undoubted harmful action, the roots being dark and 
flimsy, tips swollen and often turned upward, the greater number 
being dead. 

The plants in the solutions on the potash base line are better 
than the two sets described above. The upper parts of the roots 
of these plants are dark, the lower sections that have recently 
grown out are white and clear, but the tips are still dark and swol- 
len, and many are turned upward. 

The plants in the interior of the triangle, where all three fertilizer 
ingredients are present, are fairly good, with no great difference 
noticeable at this stage, but showing somewhat better plants near 



the center. The roots, on the other hand, are already showing 
differences. They are rather poor along the line having 8 parts 
per million of NH 3 . The upper and older part of the roots is 
dark, the newer portion is white and clear, but the tips are swollen. 
The general condition of the roots, as a whole, is a great deal 
better than in case of those in the boundary lines of the triangle. 
The condition of the roots on the line having 8 parts per million 
P 2 5 , and on the line having 8 parts per million K 2 0, are about 
the same as on the 8 parts per million NH 3 line. 

Farther within the triangle the general condition of the roots 
is much better. The solutions in cultures number 25, 32, 33, 34 
42, and 43 show by their roots that the harmful effect of the sub- 
stance has been somewhat overcome: they are lighter than the 
others, in fact most of the roots, except the upper and older sec- 
tions, are white and clear, but the tips are still slightly swollen 
and somewhat bent. 

Another property of the roots which is influenced by harmful 
bodies is that of root oxidation. It has been shown that plant 
roots possess a very appreciable power of oxidation, and that this 
power is stronger in good soils than in poor, or in their extracts 
and that harmful bodies retard this oxidation, and beneficial 
bodies augment it. Fertilizer salts were shown to increase root 
oxidation, and through this action a reduction in the quantity o 
the harmful body present was produced. 

Some further observations were made in connection with tne 
present investigation in regard to oxidation and the effect 
dihydroxystearic acid upon this function. The effect was so 
marked in the concentrations used, namely 50 parts per million, 
that the roots at the end of the experiment were found to be almos 
wholly lacking in ability to oxidize aloin used as an indicator in 
manner described in the publication referred to above. Some 
oxidizing power was still possessed by the cultures in the interior 
the triangle, where, as mentioned above, the growth was bet e 
than in other regions, although even here the power of oxidation 
was greatly impaired. Nevertheless, it is interesting to note tha 
the condition which produced the best general development 
the plants is closely associated with a power of root oxidation- 








F IG - 3- — Wheat plants growing in a culture solution containing a fertilizer mixture 
composed of phosphate 60 per cent, nitrogen 20 per cent, potash 20 per cent: 1, without 
dihydroxystearic acid; 2, with dihydroxystearic acid. 






Fig. 4.— Wheat plants growing in a culture solution containing a fertilizer mixture 
posed of phosphate 20 per cent, nitrogen 60 per cent, potash 20 per cent : I, w*' 
dihydroxystearic acid; *, with dihydroxystearic acid. 


Lower concentrations of dihydroxystearic acid would doubtless 
be better suited for a more thorough study of this matter. 

Some of the plants grown in solutions with and without dihy- 
droxystearic acid are shown in figs. 3, 4, and 5. - The cultures taken 
represent a fertilizer rich in phosphoric acid, one rich in nitrate, 
and one rich in potash. The results shown in these photographs 
are representative of the general effect over all the region of the 

triangle from which they are taken. 


The green weights obtained in the 66 cultures of the first set 
are given in the triangular diagram shown in fig. 6. It will be seen 
that, in harmony with the results given in the previous paper report- 
ing the experiment without the harmful substance, the growth 
with the single elements or along the lines where mixtures of two 
occurred was in general less than within the triangle. The region 



phosphate lines. In the dihydroxystearic acid cultures it might 


along these lines toward the nitrogen side. 

The total growth made in the 66 cultures without the dihydroxy- 




, the latter becomes 55. In other words, 
the plants with the dihydroxystearic acid made only, as an average 
of the 66 cultures, a growth of 55 per cent. It will be seen that the 


periphery of the triangle, except, perhaps, along the phosphate- 
nitrogen line. In the interior of the triangle, where the greater 



true in the region nearer to the nitrate end, in 


the dihydroxyst 







Fig. 5.— Wheat plants growing in a culture solution containing a fertilizer mixture 
composed of phosphate 20 per cent, nitrogen 20 per cent, potash 60 per cent: 1, with- 
out dihydroxystearic acid; 2, with dihydroxystearic acid. 




As has already been stated, the solutions were analyzed every 
third day for the three component fertilizer parts, phosphate, 
nitrate, and potassium, expressed as P 2 O s , NH 3 , and K 2 0. The 
original concentration in these elements was in the sum total 80 

p l«5 

Fig. 6. — Green weight of wheat grown in 66 cultures with different proportions 
of PA, NH„ and K 2 0. 


parts per million. After analysis the 
ponent parts was again calculated and the average concentration 
of these three elements was ascertained for the eight periods. 
These average concentrations will be found in the diagram in 

%• 7- 




the corresponding solution without dihydrox} 






In the diagram fig. 8 are given the original ratios of the ferti- 
lizer constituents, the ratios left in these solutions as shown by 
analysis, and the corresponding ratio of the removed constituents, 
as in the former paper. As before, the large dots in the diagram 
represent the original ratios according to the scheme previously 

P a 5 


a Fl ,°' . 7 -"~ Avera S e concentration in parts per million of P 2 O s + NH 3 + K,0 of 
the solut.on after the growth of 10 wheat plants; concentration of original solution 

Was So n n m 

was 8o p. p.m. 

explained. The circles indicate the ratio left in the solution as 
shown by analysis, and the other end of the line indicated by an 
arrow shows the corresponding ratio of the removed materials. 

In the former experiment without dihydroxy stearic acid there 
was a decided tendency for these lines to converge toward a region 
somewhat below the center; that is, the solutions near this central 
area changed least in their ratio, and the farther the ratios were 



removed from this central area originally, the more were they 
altered in the course of the experiment. This area was between the 
10 and 20 per cent phosphate line, and the area of greatest growth 

this same reg 

on. It is in this region of greatest growth, 
therefore, that the greater absorption of nutrients took place 
with the least change in ratio; in other words, the solutions repre- 

• p a°5 



Fig. 8.— Showing the ratio of the original, the final, and the ratio of the loss of 
P 2 5 , NH 3> and K 2 from the culture solution; the dots indicate the ratio of the 
constituents in the original solution; the circles show the ratio of the constituents 
in the solution after growth; and the arrows show the ratio of the decrease. 


>s for the absorption of plant nutrients. 
in fig. 8 for the experiment with dihydroxv 




will be remembered from the data already 




greatest growth under 

iditions to be nearer this end. 
In other words, when dihydroxyst 

materials removed 

solution to fall nearer to the nitrogen end of the diagram than they 
do in the case of cultures where this substance is absent. This 




Fig. 9.— Showing the ratio of the original, the final, and the ratio of the loss 


marked from the verv fir 

given in fig. 9 for the first period. Moreover, the points, for 

mstance, of the 10 per cent mixture 


lie very low, and this 

dihydroxy stearic acid is found throughout the experiment, though 

marked in all periods. The average effect ha3 

it is not equally 


The strong 




which is shown when this substance is present for a proportionately, 
and sometimes even absolutely, greater decrease in the nitrogen of 
the solutions, is strikingly shown in this first period (fig. 9). This 
varies somewhat from period to period; thus, for instance, in the 
fourth period, shown in fig. 10, the ratio change is fairly normal, 
though the usual tendency is seen again in the seventh period, a 

p a o 5 

Fig. 10. — Showing the ratio of the original, the final, and the ratio of the loss of 
*»O s , NH 3 , and K 2 from the solution, in the fourth period. 

These diagrams for the 

diagram of which is presented in fig. 11. 
several periods illustrate rather well the general tendencies brought 
out by an examination of the analytical data. As a rule, beyond 
the second or perhaps the third period the diagrammatic representa- 
tion of the result is on the whole uniform, but is influenced undoubt- 
edly by the conditions of growth during any period ; in other words, 
by weather and other conditions, which is shown perhaps quickest 

1 7 8 




The influence of light 

conditions on different days has already been discussed in the 

previous paper 

p L o 5 


Fig. ii.— Showing the ratio of the original, the final, and the ratio of the k» 









a more 

normal develor 

in the presence of dihydroxystearic acid when the culture solution* 

rnntainArl oil fU,.^ ~£ 4-U ~ £ tStl „— U„i.«~~~~ Tt waS furtu^ r 

contained all three of the fertilizer substances. It was 


apparent that there seemed to be a tendency for more 
development in a region nearer to the nitrate end. In order 



trace out this matter, a comparison of the three experiments has 
been made and is hereby presented. In this way the solutions 
which contained in all cases 50 per cent or more P 2 O s are considered 
in one group; all those which contained zo per cent or more NH, 


in a second group; and those with 50 per cent or 

a third group. By grouping the cultures thus, an average result 

mainly phosphatic, m 


mainly potassic fertilizers. 

A comparison of the average green weight in each of these 
groups of cultures was made with those in the corresponding 
groups in the cultures where dihydroxystearic acid was absent. 
The relative growths thus obtained, taking the growth without 
the dihydroxystearic acid as 100, are contained in table I. 

* • TABLE I 

Showing the average relative growth made in the group of solutions with 

AND K 2 0. 



Relative green weight 
(Green weight without dihydroxystearic acid=*ioo) 




P,O s 

50-100 per cent 

NH 3 

50-100 per cent 



K 2 

50-100 per cent 



In the first column is given the number of the experiment; in 



column the same result for the cultures of mainly nitrogenous 
fertilizers; and in the fourth 


mainly potassic fertilizers were present. The results indicate, as 
was pointed out in discussing the experiment more fully described 
m this paper, that the mainly nitrogenous fertilizers enabled the 
plants to make a more normal growth than, on the whole, do the 
other fertilizers, although with the quantities of salts and harmful 
substance used in these experiments, the plants were b 







This general relationship is even more strongly shown when the 
decreases in the concentrations of the solutions in these various 
groups are considered. The average decrease, in solutions with 
and without dihydroxy stearic acid, for each group in each experi- 
ment, is shown in table II. 


Showing the average decreases in the concentration of total constituents 
in the group of culture solutions containing fertilizer salts having 
the composition of $0 to ioo per cent of any one of the components p2o5, 
nh 3 , and k 2 0, without and with dihydroxystearic acid (original concen- 
tration = 80 parts per million). 













equals 100 

















equals 100 











equals 100 




The third column under each fertilizer group gives the relation 

between these decreases, the decrease in concentration 


dihydroxystearic acid is absent being considered as the norma 
for comparison and equal to ioo. A comparison of these relative 
effects shows that the mainly nitrogenous fertilizers give the highes 
results. In other words, the decrease with the nitrogenous ferti- 
lizers was more nearly like that observed under the normal con- 
ditions where dihydroxystearic acid was absent. This result may 
be interpreted to mean that the mainly nitrogenous fertilizers 


decreased the inhibitive effect of the dihydroxystearic acid 
although it does not show whether this is a direct or an indirect 
effect; that is, whether there is an actual decrease of this inhibits 
material or whether there is mainly the ability of the plan ^ ^ 
withstand the attacks under these conditions. Attention ^ 
already been called to the fact that this substance when in soib is 
most easily destroyed by processes which promote oxidation- an 
it should be borne in mind that the mainly nitrogenous fertile 1 
are the ones which promote the most active root oxidation by 
plants themselves. A study of the oxidation of roots in t e 


experiments showed that the dihydroxystearic • acid interfered 
greatly with oxidation, and that this action was overcome to some 
slight extent in the center of the triangle but nearer to the nitrate 
end, thus showing perhaps that there is some correlation in these 


The foregoing investigations have given the following results: 
/ i. An organic soil constituent, dihydroxystearic acid, hinders 
the growth of wheat plants, when this is present in solution in pure 
distilled water. 


2. The compound is also harmful in the presence of nutrient or 
fertilizer salts in all ratios of the fertilizer elements, P 2 5 , NH 3 , 
and K 2 0. 

3. The compound is more harmful in those ratios of fertilizer 
elements not well suited for plant growth. 

4. The harmful effect of the compound is the least in those 
ratios of fertilizer elements best suited for plant growth. 

5. The compound appears to be relatively much less harmful in 
the presence of fertilizers mainly nitrogenous than in the presence 
of fertilizers mainly phosphatic or potassic. 

6. The harmful compound modified greatly the removal of 
fertilizer elements from the solutions. The quantity of phosphate 
and potash removed was less in the presence of the compound, but 
the nitrate was not so influenced and on the whole the amount 
removed was even greater." 

7. The compound modified both amount and ratio of the three 




8. The harmful compound has the additional effect of darken- 
ing the root tips, stunting root development, causing enlarged 
root ends, which are often turned upward like fish-hooks, and 
inhibiting strongly the oxidizing power of the roots. 

9. Those fertilizer combinations which tend to increase root 
oxidation are also the combinations which overcome the harmful 
effects to the greatest extent. 

Bureau of Soils 
U.S. Department of Agriculture 

Washington, D.C. 




Chas. O. Appleman 

(with one figure) 

During the course of an investigation now in progress on the 
physiological behavior of enzymes in after-ripening of the potato 
tuber, it became necessary to investigate fully the best method for 
the quantitative determination of catalase in this organ. Some 01 
the results thus obtained may be of general interest. 


Schoenbein (1863) was the first to observe the power of various 
vegetable and animal extracts to decompose hydrogen peroxid 
with evolution of oxygen. He concluded that the enzymes occur- 
ring in the organisms were responsible for this phenomenon. -I* 115 
power of hydrogen peroxid decomposition was considered a more 
or less general property of enzymes until Loew (i) showed that 




at this time seems to be based wholly upon its sensitiveness toward 

heat, acids, and various poisons. 



In fact, its occurrence is so general that U^ 


, or any 

organ, or even a single vegetable or animal cell, that did not con- 
tain some catalase. An enzyme of such general occurrence nug 
naturally be supposed to possess an important function in t e 
economy. of nature. This may yet prove to be the case, but a 
present its position in this respect is uncertain. Little is even 


molecular instead 

atomic oxveen results from the decnrrmos;ition. and in this respe 

it differs from the oth 

Botanical Gazette, vol. 50] 



Since the decomposition of hydrogen peroxid is the most 
important property of this powerful enzyme so widely distributed, 
it was suggested by Loew that it may possess the function of 
preventing the accumulation of this toxic substance in the tissues. 
He conceived it possible and highly probable that hydrogen peroxid 
was produced in the living cells as the result of respiratory pro- 
cesses. Usher and Priestly (9) were able to demonstrate the 
presence of hydrogen peroxid during photosynthesis if the catalase 
were previously destroyed. This fact would seem to support the 
above theory, but on the other hand, Bach and Chodat (2) have 
shown that hydrogen peroxid is not a violent poison in tissues, 
since they have been able to cultivate certain plants in a medium 
containing 0.68 per cent of it. Catalase is found also in anaerobic 
organisms, a further fact which rendered the above conception 
untenable. Other authors have ascribed to catalase the function 
of protection against the peroxids of the organism, thus preventing 
injurious oxidations. It is unable, however, to decompose the 
substituted organic peroxids, such as ethyl hydroperoxid or the 
oxygenases (2). 

Probably the, most important question in connection with 
catalase at the present time is whether or not it may be considered 
as an oxidizing enzyme. It is true that it does not respond to the 
tests with the ordinary reagents for oxidizing enzymes, but this, 



enzymes is sometimes 

specific. He claims that a characteri 
produced with hydroquinone and also some with glucose and citric 
acid. Shaffer (3) thinks that this quinone-formation was 
undoubtedly due to the presence of some enzyme other than 
catalase, since he found that animal tissues always contain catalase, 
but frequently possess no power to oxidize hydroquinone. 

According to the Buchner and Meisenheimer (4) conception 
of alcoholic fermentation, the sugar is converted into lactic acid 
by the zymase, and the lactic acid in turn is split up into 
alcohol and carbon dioxid by a lactacidase enzyme. In a recent 
work Kohl (5) claims to have proved, for yeast fermentation at 
least, that catalase performs the function of the zymase in the above 


conception, and if no zymase is present the lactic acid is oxidized 
to oxalic acid by oxidizing enzymes. If zymase is present, it 
splits the salts of lactic acid up into alcohol and carbon dioxid, 
the work of the lactacidase according to Buchner and Meisen- 
heimer. This work of Kohl's, if confirmed, brings to catalase 
a very important role in physiological processes as an oxidizing 
enzyme . 


The following method was employed in all the catalase deter- 

The apparatus used i- 


shown in fig. 1. The potato was grated rapidly on a nutmeg grater 
with frequent dipping of the grated surface into calcium carbonate. 
After grating, the pulp was ground for two minutes in a mortar 
with quartz sand. The extract was then pressed lightly through 
two layers of absorbent cotton and one of cheese cloth. After 
mixing, 1 cc. was withdrawn immediately and placed into the 
bottle used for the determination, and 1 cc. of cold water added. 
The apparatus was then placed into a water bath at' 20 C. After 
the apparatus had attained the temperature of the bath, 5 cc. of 
Oakland 3 per cent hydrogen peroxid (dioxygen) were run into the 
bottle from the separating funnel 2. The stopcock a was 
opened 15 seconds before the minute. On the minute, shaking 





at the burette. Stopcock c was now opened until the menisci 
in the burettes were again level. At the three-quarter mark the 
stopcock b was opened again and closed on the minute for a reading. 
This procedure was continued with readings every 30 seconds for 







impossible to make 

— "— » w " *.c*«.aiasc in uie iresn potato exirat-i "« » — 

of the rapid degeneration during the grinding and subsequently- 

i 9 io] 






C. or below, this rapid 

may be overcome 


Fig. i. — Apparatus used for catalase determinations: 1, thin-walled bottle; 2, 
separatory funnel; 3, gas burettes; 4, thermometers; 5, weights; 6, water bath; 7, 
water meniscus; 8, water level in bath; a, stopcock for running the hydrogen peroxid 
into bottle; b, stopcock for controlling the flow of oxygen into the burette; c, stop- 
cock for equalizing the air pressure in burettes. 

without any difficulty. It is necessary, however, to maintain the 
same period of time for grinding, as there is a slight degeneration 
even under the above conditions, due to another cause, as we shall 
see later. The shakina 




carbonate. After grinding two potatoes of equal weight, one with 




and the other without calcium carbonate, they were stored at 
o° C. for 34 days. At the end of this time, 200 cc. of water were 




Ground without CaC0 3 .... 
Same extract after standing 
15 min 

Ground with CaC0 3 

Same extract after standing 
90 min 



















5 th 

Total cc. 0, 

in s MIN. 








cc. of this dilution 

and 5 cc. of hydrogen peroxid were used for the determination in 

each case. 


Effect of storage for 34 days at o° C. with and without calcium carbonate 




Ground without 


Ground with CaCO 








5 th 



Total cc. 0, 
in 3 min. 




Approximately 50 per cent of the total catalase will pass through 
an ordinary filter paper, but none through a Chamberland-Pasteur 
filter. The first fact seems to indicate the existence of an insol- 


through ordinary filter paper 

Extract of potato tuber 

Filtered through filter paper 

Portion of same extract until 










5 th 



1 . 



3 1 


Total cc. 0* 
iN3 MDf - 







(^-catalase) form. Kohl finds the 
d Loew believes it a general char- 




acter of vegetable catalase. The size of the molecule of the latter 
prevents its passage through a Chamberland-Pasteur filter. 

Table IV also shows the effect of filtering and the necessity of 
thorough mixing of the extract before taking the sample for a 
total catalase determination. 


Effect of filtering 



Extract of potato tuber 


Total cc. 2 







in 3 MIN. 




11 .9 












• • • 


2. I 


• * 



• a • • 


Supernatant liquid after settling 

After mixing, 20 min. later 

Filtered, 40 min. later 

Effect of filtering through a Ch. 

and-Pasteur filter 

Extract of potato tuber 

a) Unfiltered 

b) Filtered through Chamberland 

c) Residue of (b) made up to same 
volume as (a) 









22.8 19.3 ■|i6. 8 











14-8 13.5 

Total cc. 2 

in 3 MIN. 




Catalases from different sources show considerable variation 
in temperature relations, the point of total destruction in the cases 
reported ranging from 65 C. to 8o° C. In potato catalase, how- 

on is complete when the temperatur 
Hoff velocity coefficient for hemase 


to be 



figure applies to potato catalase, but is 

¥ io°. At 20 a destruction of the catalase 

begins, which renders the accelerating effect of higher temperatures 




destruction of the catalase, as will be seen by table VII, which also 
shows that the destruction at moderate temperatures is not due 

1 88 



to impurities in the hydrogen peroxid. Some substance in the 
potato may be freed by the grinding and brought into contact 


Joint effect of acceleration and destruction by rising 


Range of temperature 
Velocity coefficients . 

with the catalase, which effects its slow destruction. It is interest- 
ing to note in this connection, however, that Senter (6) found 
hemase to be oxidized at all temperatures above o° C. 






Potato tuber extract 

a) Exposed to 30 C. only during de- 
termination of catalase 

b) Exposed to 30 C. during exp. (a) 

then cooled to 20 C. for determina- 

c) Not exposed above 20 C 



30° c 



I St 


3d i 4 th j 5th 





cc.0 2 

in 3 


14. i 13.2 12.5 11. 9 




J3-7 *3-3 l2 -7 
14.7 13.9:13.3 



12.8 25. 




position of hydrogen peroxid. Table VIII shows not only the above 
fact, but also would seem to indicate that it is consumed in the 
reaction and that a given amount is capable of decomposing a 
definite amount of hydrogen peroxid. 



of oxygen was evolved in 5 min. Another cubic centimeter ot 
hydrogen peroxid was then added and the reaction allowed to run 
to the same point. This procedure was continued until the addi- 
tion of a cubic centimeter of hydrogen peroxid produced only 1 • 1 cc 
of oxygen in 25 min.; 6 cc. were required to bring the reaction to 




this point. A fresh portion of 5 cc. of the extract, which had stood 
at the same temperature, was now allowed to act upon 6 cc. of 
hydrogen peroxid at the same time, with the result that approxi- 
mately the same amount of oxygen was evolved as the total pro- 
duced when 6 cc. were added in successive lots of 1 cc. each. The 
experiment was repeated several times with different extracts and 
dilutions, with practically the same result in every case. 

Consumption of the catalase during the reaction with hydrogen peroxid 

Potato tuber extract 




cc. 0, 


Total cc. 0, 
evolved when 






• 5th 



5 cc. extract 





1 ■" ' ' 




# • 9 m 

5 cc. same extract 


The destruction or degeneration of the catalase would be slight 

during the time of 


above dilute neutral 

solution at 20 . Admitting a slight destruction, it would be the 
same in both cases, and would therefore not affect the general 
result. Kastle (7) suggests that catalases, like peroxidases, may 







of the peroxidase reagents, in which event we would have H 2 K0 

; 0+K0 


indicated in the above table, and with the fact that the reaction 
is decidedly an exothermal 



There is considerable evidence from animal tissues that catalase 
activity bears a relation to functional activity of the structure. 
Such a relation seems to exist in the potato, at least in respect to 
respiratory activity. Extracts from potatoes which have been 
kept for several days at o° C. show a decided decrease in catalase 
activity. Muller-Thurgau (8) has also shown that respiration 
»s greatly reduced in such potatoes. 




Catalase activity is greater at the end than at the beginning 
of the rest period. It is also greater in the large mature potatoes 


Decrease in catalase activity after storage at o° C. 

Weight of 
whole potato 

Time of 


219 gm 
214 gm 






28 days 
28 days 



12.8 10.5 





Total cc. 0, 
in 3 mix. 






3 1 



than in the small immature ones. This is true to a much greater 
extent in new potatoes than old ones, Muller-Thurgau (8) 
found that these same relations exist in regard to respiration. 



Potato at 

Wt. of potato 




Beginning of rest period 
Beginning of rest period 

End of rest period 

End of rest period 

100 gm. 
400 gm. 

147 gm. 
540 gm. 




27. 5 



19. 1 

19. 1 

Total cc. 0, 

IN 2 MLN. 






55- 6 

In table XI equal weights of morphologically similar pieces o 
new and old potatoes of about the same weight were groun l, 
washed into a volumetric flask, and made up to 250 cc. wi 
water. After thorough shaking, 5 cc. were withdrawn for t e 
test, using 5 cc. of hydrogen peroxid as before. 

Catalase activity at 

beginning and end 

OF rest per* 



Pot x TO a t 


1 \JXJ\ L\J Al 










Beginning of rest period | 
End of rest period 


21 . 2 


20. 2 


16. I 

Total cc. 0, 
in 3 ***• 


The above facts do not prove a causal relation between cata a 

and respiration, but they are highly suggestive, especially m 



light of Kohl's recent claims for catalase as an important enzyme 
in alcoholic fermentation, and therefore most likely in respiratory 


i. Comparable quantitative determinations of potato catalase 


calcium carbonate, the extract diluted immediately with ten parts 
of water to one of extract, kept at 20 C. or below, and the same 
time maintained for the grinding of the potato and the catalase 

2. An insoluble (a-catalase) and a soluble (/3-catalase) form may 
be separated by ordinary filter paper. Approximately 50 per cent 
of the total passes through. None will pass through a Chamber- 
land-Pasteur filter. 


1.5 fromo°C. to io° C. 


is an apparent progressive decrease in the velocity coefficient. 
This is due to* actual destruction of the catalase, which is not due 
in the main to impurities in the hydrogen peroxid or to oxidation 
by the hydrogen peroxid. 

4- Potato catalase is not unlimited in its power to effect the 
decomposition of hydrogen peroxid. It seems to be consumed 
in the reaction and a given amount is capable of decomposing a 
definite amount of hydrogen peroxid. 

5. The catalase activity bears a relation to the respiratory 
activities in the potato, decreasing under the same conditions as 
respiration. • - 

I wish to acknowledge my indebtedness to Dr. Willi 
Crocker, under whose supervision this work was pursued. 



i- Loew, O., Catalase. U.S. Dept. Agr., Rep. 68. 1901. 
2 - Bach, A., und Chodat, R., Ueber den gegenwartigen Stand der Lehre 
pflanzlichen Oxydationsfermente. Biochem. Centralbl. 1:417-421. 1903. 

3- Shaffer 

Physiol. 14:299. 1905. 



4. Buchner, E., und Meisenheimer, J., Ueber die Milchsauregahrung. 
Liebig's Ann. 349:125-140. 1906. 

5. Kohl, F. G., Ueber das wesender Alkoholgahrung. Beih. Bot. Centralbl. 
25:115-126. 1910. 

6. Senter, G., Wasserstoffsuperoxydzersetzende Enzyme des Blutes: L 
Zeit. Physikal. Chem. 44:257-263. 1903; 51:673-705. 1905. 

7- Kastle, J. H., The oxidases. Public Health and Marine Hospital 
Serv., U.S. Hygienic Lab. Bull. 59:139-140. 1900. 

8. Muller-Thurgau, Herman, Beitrage zur Erklarung der Ruheperioden 
der Pflanzen. Landwirtsch. Jahrb. 14:859-863. 1885. 

9. Usher, F., and Priestly, J. H., A study of the mechanism of carbon 
assimilation in green plants. Proc. Roy. Soc. London B 77 : 3 6 9"37 6 - 
1905; 78:318-327. 1906. 

10. Gruss, J., Enzymwirkungen aus Wundrant der Kartoffknolle. Zeit. 
Pflanzenkrankheiten 17:69-70. 1907. 



Frank M. Andrews 

Twin hybrids are produced according to DeVries "by the com- 
binations Oenothera biennisXO. Lamarckiana, and O. muricataXO. 
Lamarckiana, and by those of some of their derivatives/' 1 Besides 
laeta and velutina I shall describe here O. Lamarckiana and O. 
biennis. O. biennis, along with some other species, was introduced 
into Europe a long time ago. The origin of O. Lamarckiana, or 
large-flowered evening primrose, is involved in obscurity. In an old 
potato field near Hilversum, in the year 1886, DeVries 2 found a 
great many specimens of O. Lamarckiana which were observed to be 
in a state of mutability. I have visited this primrose field to observe 
and collect specimens. There are comparatively few of them left, due 
to the fact that they are rapidly being crowded out by pine trees that 
have been planted there. It is apparently only a question of a little 
time, unless something intervenes, until they will have disappeared 
entirely from that locality. O. Lamarckiana produced new forms by 
mutation, not only in the wild state in the field near Hilversum, but 
also in cultivation. This latter fact was proved by transferring rosettes 
from the field to the garden, where the plants could be protected and 
watched throughout their entire existence. This method, together with 
that of sowing the seeds 3 of these plants under controlled conditions, 
has allowed quite a number of mutants to assert themselves and to be 
recognized. Unless such methods as were introduced by DeVries 
in his garden at Amsterdam are followed, the detection of a mutation 
is much more apt to be only accidental. Furthermore, these methods 
when used for rapid multiplication in the garden under controlled and 
protected conditions may also be conducive to an increase of mutants, 
and hence their more certain and frequent recognition. For "such a 
rapid multiplication in the course of a comparatively few years" (as 

1 DeVries, Hugo, On twin hybrids. Bot. Gazette 44:401-407. 1907. 


DeVries, Hugo, Mutationstheorie 1 : 152, 401-407. 
3 De\ries, Hugo, Species and varieties, p. 525. 1905. 

[Botanical Gazette, vol. 50 


was noticed with O. Lamarckiana near Hilversum) "may perhaps be/' 
says DeVries, "one of the conditions for the appearance of a mutable 
period. " 4 The number of new forms that have arisen and been 
detected by DeVries in the genus Oenothera since his discovery 


of the mutating O. Lamarckiana at Hilversum in 1886 is already 

The following description of the forms O. Lamarckiana, O. biennis, 
laeta, and velutina represents in each case the average state of affairs 
as shown by a great number of examined specimens. 

O. Lamarckiana is an erect, strong plant. Its stem is cylindrical. 
1-1 . 5 cm. in diameter, 1 . 5 or more meters in height, and sparsely 
covered with coarse hairs. Some of the hairs on the stems are glandu- 
lar. The main stem is more or less branched at the top, and also has 
a tendency to produce smaller branches at the bottom when it grows 
in an open situation. All of the secondary branches may produce 

The leaves are somewhat broadly lanceolate, dark green, their 
margins distinctly dentate, and the lower ones strongly petioled. 
Both the upper and the lower surfaces, especially in the adult forms- 
are conspicuously undulated or wrinkled, and only slightly pubescent. 

In order to get buds showing as great uniformity as possible, the} 
were chosen just before they were ready to open. The buds of 0. 
Lamarckiana are the largest of any of the three forms mentione . 
being about 5 . 5 cm. in length, about 1 cm. broad near the base, a 
tapering slowly upward to a width of about 2 mm. at the top. They 
are slightly pubescent. This pubescence of the buds of all the three 
forms studied consists of two kinds of hairs, a small, thin-walled- 
cylindrical hair, and a larger, thick-walled, pointed one. In 0. la- 
marckiana both of these hairs are much larger than in any of t e 
other forms of Oenothera to be described, and especially is ** 
true of the pointed hairs. The buds are almost circular in cross- 


ed rather 

The flowers are large and bright yellow, and arranged rat er 
closely together in a broad spike. Each flower is subtended by a 
sessile bract and stands singly. The ovary is partly glandular- 
pubescent, and the four lanceolate sepals are strongly reflexed, gen 

4 Df\ "ries, Hugo, Mutationsth* rie i:im. 


erally more or less united at the top, and somewhat pubescent. 
The calyx tube is a little longer than the sepals. The cell walls of 
the epidermis of the sepals of O. Lamarckiana, especially the inner 
epidermis, are very irregular and zigzag in form. At the angles of 
these zigzag places the walls of the inner epidermis are marked by 
conspicuous thickenings. In the case of the outer epidermis the 
thickenings at these places extend away from the main wall as 
T-shaped processes. The same is true, but to a slightly less degree, 
for the inner and outer epidermis of the petals. The four very large, 
broadly obovate, entire petals are very conspicuous, and almost or 
quite equal the length of the calyx tube. One of the distinguishing 
features of O. Lamar ckiana is the tall stigma, which far exceeds 
in height the stamens, and on this account self-pollination can- 
not occur. The anthers of O. Lamarckiana open in damp or 
rainy weather as well as on clear days, but this, as we shall see 
later, is not true of some forms, as for example the twin hybrids. 
This can b? accomplished in the parent species, O. Lamarckiana, 
because at maturity the tissues of the cell walls of the anther are 
completely differentiated and become free, so that the outer wall may 
open out in the usual way. 

The capsules are four-celled and open by as many valves at 
maturity. The pubescence on the capsules is not so dense as on the 
ovary. As the old flowers disappear and new ones appear, long 
rows of capsules are left on the older parts of the spike. 

Oenothera biennis is not as stout as 0. Lamarckiana. Its stem is 
erect, cylindrical, branching, and sparsely covered with coarse but 
non-glandular hairs. It is branched, is about 1 
1-1.5 meters in he : ght. 

The leaves are dark green, broad (somewhat more so than in 
0. Lamarckiana), about three times as long as wide, with margins 
slightly wavy, and all of the stem leaves are petioled. In the last 
two respects the Holland species is somewhat different from the more 
narrow-leaved American species of O. biennis, which has only the 
"lowest petioled" 5 and the margins "repand-denticulate." 5 These 
are among the differences that show that the form O. biennis, which 
is very common in the United States, is not the same as the O. biennis 

» Brittox and Brown-, Flora of the northern U.S. and Canada 2:4*6. 



of Europe. The upper and lower surfaces of the lower leaves are 

* 1 1 « . 



The buds are much smaller than those of O. Lamarckiana, aver- 
aging i • 5 cm - long, 6 mm. wide at the base, and i . 5-2 mm. wide at the 
top. Besides being smaller, they are different in form from those 

One side of the bud is rather straight, and the 
other more or less gibbous. Comparing this with the American species, 
as shown by Britton and Brown's figure, 6 there is another dif- 
ference, for in the American forms the buds are larger and more 
pointed, especially at the apex, than in the Holland type. The hairs 
on the buds are only about half the size of those of O. Lamarckiana. 

The flowers are much smaller than those of O. Lamarckiana and 
are bright yellow. Each flower stands in the angle of a nearly sessile 
bract on the rather narrow spike. The calyx tube in this form 
is 1 . 5-2 times the length of the reflexed sepals. The density of 
the pubescence on the sepals is about the same as for O. Lamarckiana. 
The irregularity shown in the epidermal cell walls of the sepals is 
much less in O. biennis than in O. Lamarckiana, and especially is 
this true of the inner epidermal cell walls, which are only wavy, and 
in which the thickenings are almost completely absent. . The cells 
of the outer epidermis are also decidedly smaller and have but 
few thickened places. 

The four petals are broad, slightly unequally lobed, somewhat 
indented at the apex, and larger than the sepals. The cells both of 
the inner and outer epidermis are smaller, more regular in form, and 
are, especially those of the inner epidermis, almost devoid of the 
angular thickenings shown by O. Lamarckiana. 

One of the chief differences in the flowers of O. Lamarckiana and 
O. biennis is that in the latter the stamens are of the same height 
as the stigma, and the anthers are in contact with the stigma 
in the bud. Therefore self-pollination nearly always results, whereas 
in O. Lamarckiana it was generally brought about by insects. The 
four anther cells do not always open simultaneously, but some 
may have partly opened and shed their pollen before the opening 
process in the other cells has begun. The ovary is four-celled and 

« Britton and Brou v, p. cit. 2 : 486. Jig. 2 579 . 


The capsules are almost pressed against the stem. They are 
about 24 mm. long, somewhat pubescent, and remain as the flowers 
disappear in long rows on the spike, as in O. Lamarckiana. These 
capsules of O. biennis, according to Brixton and Brown's figure, 7 
seem to be differently curved and more pointed toward the top. 
Instead of the usual number of parts of the flower of O. biennis, I 
have observed some where there were three sepals, three petals, six 
stamens, and three stigmas. Deviations of a similar kind I have 
noticed in other plants. 8 

The stem of laeta is branched, about the same height as that of O. 
biennis, and almost intermediate between the two parents as to 
stoutness and pubescence. 

The leaves are broad, bright green, and while wrinkled they are 
hardly as much so as in O. Lamarckiana. They are considerably 
more pubescent than those of O. biennis and slightly more so than 
those of O. Lamarckiana. 

The buds differ in size and form from those of the other forms. 
They are more pubescent than in O. Lamarckiana or O. biennis^ 
smaller than in O. Lamarckiana, and larger than in O. biennis. In 
cross-section they are almost round. 

The hairs of the buds and other parts of the plant are distinctive 
as to size, for they are considerably longer than those of either O. 
biennis or velutina, but smaller than those of O. Lamarckiana. 

The lanceolate and strongly reflexed sepals are about three-fourths 
of the length of the calyx tube, and are somewhat more pubescent 
than in any of the preceding forms. 

Both the form and the size of the cells of the outer and inner epi- 
dermis of the sepals, as well as of the petals, of laeta are different 
from those of either of the three other forms mentioned here. 

The relative size of the stamens and style is another distinctive 
feature, for the style is somewhat higher than the stamens, being 
less pronounced than in O. Lamarckiana. Both the stamens and the 
style are much smaller than in O. Lamarckiana. The stamens about 
equal in size those of O. biennis and velutifta. The anther cells 
behave differently from any of the previously described forms, in 

i Brittox and Brown, op. cit. 2:486. fig. 257Q. 

8 See my paper in Proc. Indiana Acad. Sci. for 1905 and 1906. 


that in damp or rainy weather they do not open at all, and some- 
times very imperfectly even on sunny days. I have observed anthers, 
one or two of whose cells never open, and the remaining cells do 
so only partly. The anthers seem to develop normally in every 
respect except that the tissues do not become differentiated com- 
pletely at the usual point of opening, so as to allow the escape 
of the abundantly formed pollen grains in the ordinary way. It 
was often necessary to exert strong pressure on the cover glass before 
the anther wall would separate at the usual place, and this was true 
even in thin cross-sections. 

The ovary of laeta is more pubescent than that of O. Lamarckiana 
or O. biennis. 

The capsule is about the size of that of O. Lamarckiana, but 
different in form from either parent. They form very long spikes 
of fruit before the close of the flowering period. 

Velutina, the other twin hybrid form, is also distinctly different 
from laeta and all other forms. 

Its stem is about the same height as that of laeta, somewhat more 
branched, and light grayish, due to the dense pubescence. The 
hairiness of the whole plant is much more pronounced than in any 
of the other forms described, and this fact alone would serve to dis- 
tinguish it from any of them. The stem is not quite so stout as 
of laeta, and the aspect of the whole plant is somewhat less vigorous. 

The leaves are grayish green, narrow (more so than in laeta)- 
trough-shaped, tapering toward each end, and much more pubescent 
than the leaves of any of the other forms. 

The buds are shorter than those of laeta, but have a greater 
diameter and are much more hairy. The hairs from the buds, as 
in the other types, are of two forms, cylindrical and pointed. The 
cylindrical hairs are only a little larger than those of O. biennis, 
but smaller than in the other types. The pointed hairs are exceeded 


Lamar ckia 

O. biennis and laeta. 


those of laeta and all the other forms in another respect. In all n 
other forms the pointed hairs are smooth, but in velutina they are 
densely covered by numerous and conspicuous elevations. 
The flowers are somewhat smaller than those of laeta. 


The sepals are only about one-half the length of the calyx tube, 
whereas in laeta they are about three- fourths of its length. This 
difference is very evident on comparison. The sepals of velutina are 
also much more hairy than any of the other forms. 

The cells of the outside epidermis of the sepals of velutina are 
about uniform in size with those of laeta, but the walls are much 
stronger. The cells of the inner epidermis are more uniform in size 
than in laeta. 

The petals are more deeply indented at the top than in laeta, are 
somewhat smaller, and light yellow. The cells of the inner and 
outer epidermis are more nearly isodiametric than in laeta, and their 
cell walls are much more zigzag, and thickened at the angles. 

The stamens are the same length as the stigmas. In this respect 
they differ strikingly from laeta and O. Lamar ckiana, but resemble 
0. biennis. As in laeta, they open some of their anther cells at 
least, on sunny days, but on damp and rainy days they remain closed. 
I have noticed instances, as in laeta, where some of the anther cells 
never open, no matter how ideal the conditions. I have found it 
necessary to use considerable force in order to cause even thin cross- 
sections of the anthers to open their cells. This, as in laeta, is due 
to the fact that the tissues at the point of opening never separate, but 
remain more or less completely grown together. This non- opening 
of the anthers of Oenothera forms, however, is not confined to those 
mentioned above. I have visited another primrose garden at Haar- 
lem which is conducted by Mr. A. R. Schouten and Mr. J. Jeswiet. 
In this garden are grown, among other forms, O. scintillans and O. 
gigas, and Mr. Jeswiet kindly informed me that they often find it 
necessary to open the anthers with a needle in order to free the pollen. 
Much of this pollen, as in our twin hybrids, is not sterile, but in some 
cases would never escape naturally from the anthers. Mr. Jeswiet 


types they have at Haarlem 

0. scintillans must be opened with a needle to obtain the pollen, 
and also many of the anthers of the form O. gigas. 


0* pubescence. 

The capsule resembles that of laeta, but is curved near the top. It 
is strikingly different from laeta in regard to its denser pubescence. 



One further difference remains to be recorded in reference to the 
petals of the flowers of O. Lamar ckiana, O. biennis, laeta, and vehtina. 
As we look at these flowers on their respective plants, the smaller 
and even the more striking differences of the petals are apt to escape 
notice. These can be seen most clearly by removing a flower of each 
of the forms just named and spreading them out carefully on white 
paper. To do this best and to fasten them to the paper, a coat of 

"age is first applied. This allows the petals to be arranged 
without injury, and at once discloses their true size, form, and 
relation to each other. 

T In O. Lamarckiana the four large petals do not touch one another 
even at their nearest edges. Their narrowed bases leave wedge- 
shaped openings. The lobes of the petals are equal. 

In O. biennis the form of the petals is at once seen to be different, 
and they do not touch each other. The petals do not run down to 
the base as straight lines, but curve in so as to leave a nearly flask- 
shaped opening. Furthermore, the petals are slightly unequally lobed, 
but always in one direction. 

Laeta is very plainly different from velutina by the conspicuous 
and considerable overlapping of its four petals. These overlapped 
edges of the petals leave inclosed spaces at their base. These petals 
are also seen to be considerably larger than those of velutina. 

The petals of velutina resemble those of O. biennis in general 
form, but are somewhat unequally lobed and larger. The spaces 
between the petals of velutina are larger and more nearly closed by 
the broad part of the petal than in O. biennis. 

The cross-sections of the stamens mentioned in this paper were 
made as follows: Holes were made in cork with a small cork-borer 

the anthers were placed carefully in them, and the holes were then 
filled with liquid glycerin jelly. After the glycerin had again become 
firm, the piece of cork with the specimens was placed in 96 per cent 
alcohol and allowed to dehydrate and further solidify the glycerin, 
which required two to three days. The sections were then made 
through both cork and specimens free-hand with a razor, and mounted 
for study in glycerin jelly. Such specimens, of course, can be made 
permanent by sealing the cover glass with Canada balsam. 





anthers. The anthers open both on rainy and sunny days, whereas 
some of the other forms open their anthers only on sunny days. 

2. The European species, O. biennis, differs from the American 
species. Its anthers and stigma are of the same height, instead of 
being of different height as in O. Lamarckiana. Its petals, pubes- 
cence, and most of its cells are the smallest of any of the forms 
described, while those of O. Lamarckiana tend to be the largest. 

3. The twin hybrids laeta and velutina show themselves by their 


foliage, flowers, the greater density and character of the pubescence 
in velutina, as well as the arrangement and form of the cells, to be 
distinct, and, in so far as has yet been investigated, constant forms. 

4. On rainy days the anthers of the twin hybrids do not open, but 
only on sunny days, and then only part of them, whereas in all the 


This non- 

opening of the anther cells may probably be the beginning of 

In conclusion I take pleasure in expressing my thanks to Professor 
DeVries for having placed at my disposal one of his laboratories 
and material from his garden for this investigation, and also for 
his many kind suggestions and assistance. 

University of Amsterdam 



Edward W. Berry 

(with two figures) 

Eocene plants have been thus far unknown along the Atlantic 
border, the most easterly known flora of this age having been that 
of the so-called Eolignitic of the Mississippi embayment, which 
while abundant is for the most part uncollected and undescribed. 
Contrasted with this paucity of eocene plants in the east, the 
western interior Eocene has a flora which in number of species 
probably exceeds the sum total of all the later tertiary floras of 
North America. There are also a number of eocene floras along 
the Pacific coast and in the far north as far as Alaska, the mouth 
of the Mackenzie River, and Greenland. 

The following brief paper is concerned with a most interesting 
eocene flora of Claiborne age recently discovered, and studied b> 
the writer in connection with the cooperative investigation ot the 
coastal plain in charge of Dr. T. Wayland Vaughan of the U.S. 
Geological Survey. These notes partake of the nature of an 
abstract of this study, which will eventually be published in full 
by that organization. 

The localities are all in eastern Georgia, where there is a markec 
transgression of the Claiborne sediments, burying all traces o 
Upper Cretaceous or earlier Eocene deposits and coming to res 
upon beds of Lower Cretaceous age or even in some instances up 
the crystalline rocks of the eastern Piedmont. The bulk of t e 
plants come from the vicinity of Grovetown, about seventeen 
miles west of Augusta near the southeastern border of Colum 
County. In this area the deposits, which consist for the m 
part of porous laminated light-colored clays alternating with heavy 
beds of sand, and more rarely beds of lignite, occupy a pre-Claibor 
estuary eroded in the Lower Cretaceous and outcropping as a narr 




Baniotcal Gazette, vol.50] 

[* c 


main body of the Claiborne deposits in Richmond County in a 
northwesterly direction for a distance of about eighteen miles. 

The plants are intimately associated with a few estuarine and 
shallow water marine invertebrates, such as Modiolus, Ostrea, 
Nucula, Leda, Cytherea, etc., which fully corroborate the age of the 
deposits derived from the study of the flora, where the data for 
correlation, in the absence of known Claiborne floras, consisted of 
comparisons with geographically remote floras like those of the 
Green River beds of Wyoming and those of the Paris basin and 
southern England. There are certain features, however, notably 
a marked indication of a rise in temperatures and increased humid- 
ity, which, in so far as they are known, characterize middle eocene 
floras everywhere. This is known to be the case in America, it 
is markedly shown in the floras of England and France, and is 
emphasized in the numerous late eocene floras from a large number 
of arctic localities. 

Among the forms collected from the Georgia Eocene are new 
species of Acrostichum, Arundo, Castanea, Conocarpus, Dodonaea, 
Ficus, Malapoenna, Momisia, Pisonia, Potamogeton, Rhizophora, 
Sapindus, Terminalia, and Thrinax, including the first fossil occur- 
rence of species of Conocarpus, Momisia, and Thrinax. 

The foregoing forms represent twelve families and include one 
fern, three monocotyledons, and ten dicotyledons. No gymno- 
sperms, which are usually represented in European Lutetien floras 
by at least the genus Podocarpus, have been discovered. It will 
be observed that only two forms, the Castanea and the Potamogeton, 
are not coastal forms, and the latter is an aquatic whose presence 
associated with coastal swamp or strand plants is not difficult to 
explain. The Castanea then apparently represents the only upland 
fype preserved in this flora, and as it is not common, the presump- 
tion is strong that it was brought down to the basin of sedimenta- 
tion by some eocene river, most likely by the river which originally 
occupied the trough of the subsequent estuary. 

Turning to the remaining twelve species, we may enumerate 
^vith profit their closest allies in the existing flora. The Acrostichum 
is represented by A. aureum Linn., the Arundo by A. Donax Linn., 
the Conocarpus by C. erectus Linn., the Dodonaea by D. viscosa Linn., 




the Ficus by various tropical American figs, the Malapoenna by 
M. geniculata (Walt.) Coulter, the Momisia by M. aculeata (Sw.) 
Kl., the Pisonia by P. macranocarpa Donnell Smith, the Rhizophora 
by R. Mangle Linn., the Sapindus by S. saponaria Linn., the 
Terminalia by T. phaeocarpa Eichler, and the Thrinax by various 
West Indian species of that genus. 

When we consider the habitat of these modern forms and that 
of their allies, we find that they inhabit the tidal nipa swamps of 
the orient, the mangrove swamps of the orient and the Occident, 
the beach jungle of the strand, or the landward side of coastal 
sand dunes in the tropics. Nearly every one of the fossil species 
is represented by forms found in the existing flora of the Florida 
keys or along the shores of peninsular Florida, some like Conocarpus 
flourishing equally well on either muddy or sandy shores. Every 
species (except Castanea) is represented in the American tropics 
and four of these existing representatives (Conocarpus, Dodonaea, 
Rhizophora, and Sapindus) range northward to Bermuda in the 
path of the gulf stream. In looking over Schimper's classical 
Indo-malayan strand flora, the following forms, which are strict!} 
comparable to the Georgia eocene plants, were noted as being 
more or less prominent elements in the oriental strand flora. 
Acrostichum {Chry sodium, i sp.), Dodonaea (i sp.), Eugenia^ (2 
species which are represented in the west African and American 
tropics by the allied genus Conocarpus), Ficus (1 sp.), Malapoenna 
(Litsaea, 1 sp.), Pisonia (4 sp.), Rhizophora (2 sp.), Sapindus (1 sp-, 
and Terminalia (1 sp.). Of these forms the Sapindus, Terminals 
Dodonaea, Ficus, Malapoenna, and Pisonia are more particular} 
elements of the littoral forest (beach jungle of Kurz, Barringtoma 
formation of Schimper), while the others are integral members or 
rather intimately associated with the mangrove or the nipa associ- 
ations. It is really remarkable to what an extent the identitie 
elements in the Claiborne flora corroborate one another and de - 

t nP 

nitely denote the character of their habitats, which were, on 
one hand, mangrove swamps at certain points where the condition 
were favorable, and elsewhere the vegetation of the rain fores 
which clothed the sandy beaches or was developed behind dunes, 
which possibly formed the highest inner margin of the beach inpla ces 




Certain rather definite climatic deductions appear to be justi- 
liable from the foregoing facts. None of the modern representa- 
tives of this eocene strand flora flourish north of the winter iso- 
therm of about 50 F., and the majority do not occur north of the 
winter isotherm of 6o° F. None of the fossil forms, except possibly 
the Potamogeton, the modern species of which range over a great 
many degrees of latitude, or the Castanea, which likewise has a wide 
range, would be expected to occur outside of the areas where the 
modern subtropical rain forests are developed. We would expect 
the Claiborne climate in the area under discussion, at least at low 
elevations along the coast and in proximity to the eocene gulf 
stream, to have been uniformly humid with an abundant and 
evenly distributed rainfall. The temperature would have been 
uniform, not necessarily extremely hot, and any degree of winter 
cold would have been fatal. 

These considerations are in a large measure corroborated by 
what we find to have been the conditions in Europe at this time. 
It is well known that the middle eocene floras of Europe show 
many tropical characters absent in the earlier Eocene. These 
first become marked in the fruits from the London clays and the 
leaves from Alum Bay and in homotaxial deposits on the continent, 
and while it was once the fashion to see Australian affinities in 
these floras, they show closer affinities with the modern floras of 
Malaysia and tropical America. The accompanying sketch map 
(fig. 1) will bring this out very well. It shows the area of distribu- 
tion of the modern genus Nipa and the eocene genus Nipadites, 
which is indistinguishable from the modern Nipa. The latter has 
a single species inhabiting the tidal waters of the Indian Ocean, 


vying with the mangrov 


in both form and structure to the fossil fruits which form the basis 
for the genus Nipadites. As the map shows, these tropical or 
subtropical floras ranged northward in Europe at this time to 



habitat for one or more species of Acrostic hum closely allied to that 




of Georgia, and other forms are not wanting. In the Paris basin a 
species of Pandanus is associated with the Nipadites, while at 
Bournemouth Gleichenia and various tropical Polypodiaceae occur. 
The Lutetien stage of the Eocene in Europe and the Claiborne 
stage in America are both characterized by a remarkable trans- 
gression of the sea, the approximate shore line of the Claiborne sea 
being shown on the accompanying map (fig. 2). This map makes 
no pretence to exactness, which would be impossible on so small 


Sketch map showing geographical distribution of the existing genu 
hi pa Thunberg and the eocene genus Nipadites Bowerbank; horizontally lined area 
shows single existing species; vertically lined area shows eocene species. 

a scale map, even if the areal extent of the Claiborne deposits were 
accurately known. What it is intended to illustrate is the relation 
of land to sea and the line of advance of the flora which was neces- 




the whole of the 

In studying the Georgia flora, the writer became much intereste 


far as they are known are worthy of enumeration. It has been 
shown that certain species of Eugenia. Terminalia. Rhizophor< 





Pisonia, and Sapindus have become adapted for dispersal through 
the agency of ocean currents, by specialization of their fruits or 
seeds, which have developed air chambers or woody husks for 
buoyancy, and practically impervious seed coverings for the 
exclusion of sea water from their vital parts. Everyone is familiar 



Sketch map showing the approximate shore line of the middle eocene 
sea in southeastern North America. 

with the extreme specialization of the mangrove, which sends its 
germinating plantlets out into the world fully prepared to anchor 
themselves in water of the required depth, but the 
specializations of the other members of these floras, though less in 
degree, are almost as effective, judging by the present geographical 
distribution of these genera, so that ocean currents must play a con- 
siderable role in distribution, despite the contrary opinion of De 



Candolle. Other members of the Claiborne flora may be sup- 
posed to have been distributed by fruit-eating birds. This seems 
clearly to apply to Ficus and probably to Malapoenna. 

It can be readily shown that the existing flora of peninsular 
Florida, the Bahamas, Bermuda, etc., contains a large element 
which has been derived in comparatively recent geological times 
from the south. In the case of the Bahamas and Bermuda, almost 
their entire flora has had such an origin. If, however, we study 
geographical distribution in the light of historical geology, we find 



;, if not earlier. Experience teaches 
us that nearly all modern plant families, unless it be the most 
specialized forms like the orchids among the monocotyledons or 
the composites and their allies among the' dicotyledons, were at 
some time more widely, distributed than they are at present, and 
that the details of modern geographical distribution represent in a 
less degree the interchange of types between different areas than 
they do the greater or less degree of segregation of descendants 
of forms once spread over much wider areas. Uniformity appears 
to have been the rule during geologic history and not the exception. 
From a study of the Claiborne flora it is evident that the main 
elements of the modern flora of tropical America reached as far 
northward in the Eocene as latitude 33 and probably much farther 



from a study of the tertiary faunas of Florida, places a marked 
change in climate at the close of the Oligocene, and accounts for 
it by the elevation in the Florida area and the shifting to the east- 
ward of the gulf stream with an inshore southerly flowing cold 


Thus while the strictly modern movement of the subtropical 


northward as the various coral islands of the Bahamas became 




The Johns Hopkins University 

Baltimore, Md. 

* Dall, Trans. Wagner Inst. 3:1549, 1550. 1903. 



David M. Mother 

During the past two or three years the writer has been collecting 
data from a study upon the gametophyte of Onoclea Struthi- 
opteris with reference especially to the dioecious character of the 
prothallia, and while this study is in no way completed, it has been 
deemed advisable to make public some of the facts observed, 
inasmuch as the data obtained supplement those set forth in a very 
interesting paper, appearing in a recent number of this journal, by 
Miss Wuist. 1 

The impression is general that the prothallia of Onoclea Strut hi- 
opteris are dioecious, 2 and although this is generally true, it is not 
strictly so, as bisexual prothallia may readily be found under 

cultural Conditions. In 1907, in 




The writer has recently begun the study of a fern, whose prothallia have 
been reported as strictly dioecious, and if the spores of the same are well nour- 
ished, female prothallia will predominate, while with poor nourishment the 
vast majority of spores will give rise to male gametophytes. An examination 
of cultures grown under favorable conditions for laboratory use, in which the 
spores were sown thickly, showed that certain spores produced strictly male 
plants, others female, and still others bisexual prothallia. A small number of 
spores were isolated and grown under similar and very favorable conditions, 
with similar results. The pure males were almost equal in number to those 
bearing the female organs, while the bisexual plants were few, being about 
4 per cent of the whole number. The foregoing results seem to lend encour- 
agement to the view that environmental conditions may have much less to do 
with the development of male and female prothallia than had hitherto been 
supposed. The very brief study showed clearly that in the fern in question 

1 Wuist, Elizabeth Dorothy, The physiological conditions for the development 

of monoecious prothallia in Onoclea Struthiopteris. Bot. Gazette 49:216-219. 

2 Campbell, D. H., Mosses and ferns. Edition 2, p. 314. 1905. 

3 Mottier, D. M., History and control of sex. Proc. Ind. Acad. Sci. 1907. pp. 

20 9 ] 

[Botanical Gazette, vol. 5° 


210 BOTANICAL GAZETTE ' [September 

there is a great mortality among the spores, which, as can be readily seen, 
vary greatly in size. Among the first things to establish in this and similar 
cases, is whether mortality is greatest among the smaller or larger spores, and 
whether the prothallia springing from the smaller spores tend to remain small 
and produce only antheridia, while the larger female plants arise only from the 
larger spores, and so on. 

The fern in question was Onoclea StriithiopteriSj although the 


name of the plant was not stated in the address in question. 

Miss Wuist {op. cit. p. 217) states that about 1 per cent of the 

prothallia of soil cultures were monoecious. This estimate seems 

to me to be much too low, inasmuch as an examination of a larger 

number of prothallia, since the publication of the paragraph just 



from which this estimate was made 

soil under good cultural conditions. The plants, although grown 
thickly, were vigorous, normal in every way, and in fact the 
cultures seemed to leave nothing to be desired. The monoecious 
prothallia were as a rule young, bearing one to several archegonia. 

in diameter. Antheridia were very rarely 
found on older and larger prothallia, although little attention 
was paid to exact size and measurement. The antheridia were 

m m 


upon the older tissue; very rarely did they appear at the margin 
of the prothallium. In one case an antheridium was found upon 



I did not 

rely at all upon an examination of fresh material, but all plants 
were carefully fixed, stained in toio with borax carmine, cleared, 
and examined in clove oil or balsam. In this way a single srnal 
antheridium could easily be found upon a prothallium, and cast 

were not infrequent of a single antheridium appearing inconspi<-« 
ously among the rhizoids. It has been my experience that one or 
two immature antheridia among the rhizoids are easily overlooke 
when the plants are examined alive. In material thus stained an 
cleared, one may also more readily and accurately detect s i 
small male plants adhering among the rhizoids of the larger pro- 
thallia bearing archegonia. 

In order to ascertain whether crowding was in any way re»po 


sible for the sex of the gametophytes, cultures were made by sowing 
spores thinly upon the soil and keeping the cultures under the most 
favorable conditions available. One such culture may be men- 
tioned in detail. In this 207 spores were sown and 150 prothallia 
harvested. Of these 78 were pure males, while 72 bore archegonia, 
and of the latter 10 per cent were monoecious. Cultures were 
also made to determine, if possible, whether the spores of any 
given sporangium were male or female, as has been reported 
for certain liverworts. In this set of experiments the spores of 
individual sporangia were sown upon earth in separate dishes, with 
results similar to those from spores sown from many sporangia, 
that is, both male and female prothallia were grown; of the female, 
those bearing archegonia, some few were bisexual. Such experi- 
ments show conclusively that sporangia do not bear spores giving 
rise exclusively to male or female prothallia, and the fact that 
prothallia are bisexual is proof that no sex-differentiating chromo- 
some is present in this fern. 

No effort has been made up to this time by the writer to induce 
the development of antheridia upon "female" prothallia. Inter- 
esting experiments along this line have been carried out by Miss 
Wuist, as mentioned in the foregoing, with the following results: 
monoecious prothallia were obtained by transferring the plants 
from distilled water to Knop's solution, in which they were further 
cultivated; by transferring "female" prothallia from soil to a 
nutritive solution, and by transferring from one nutritive solution 
to another. 

These experiments indicate that gametophytes bearing arche- 
gonia only may be induced to develop antheridia as a response to 
environmental conditions. Knowing that dioecism has not been 
completely established, these results, though important, are not 
surprising. They do not show a very far-reaching influence of 
environmental factors in regard to the regulation of the develop- 
ment of sex cells ; for I have frequently observed in older cultures, 
that is, about five months after sowing, that certain "female 
prothallia, in which an egg failed to be fertilized, developed from 
the oldest parts numerous small lobes which give that end of the 
prothallium a finely fringed appearance, and upon these lobes 



numerous antheridia were produced. Such prothallia were well 
nourished, with a deep-green color, and continued to develop the 
archegonial meristem, which produced continuously numerous 
archegonia. The specimens before me at this writing are 5-6 mm. 
in length. As stated, the antheridial lobes were confined generally 
to the posterior or older parts of the prothallia, but they may 
extend a short distance forward along the margins. Antheridia 
were not observed, except in very rare cases, upon the archegonial 



of the opposite sex, either normally or in response to conditions 
controlled in a measure by the experimenter, may readily find a 
satisfactory explanation, but the case may be different in regard to 
the prothallium that produces only antheridia. If purely male 
prothallia of Onoclea Struthiopteris are merely those that do not 
become large enough, because of nourishment, to bear archegonia, 


to develop the female organ? The writer knows of no case on 
record in which this has been done. If the male plants could be 
made to develop the archegonial meristem and its archegonia, 
then it may be said that the environment (which of coures means 
chiefly nutrition) controls sex to that degree. The writer has not 
been able to believe that male prothallia of the fern in question 
are merely those that do not become large enough to bear arche- 
gonia, but that certain spores are predetermined to develop into 
purely antheridial plants, and others into those capable of form- 
ing archegonia. If there were a sex-determining chromosome, 
then one half of the spores would be male and one half female, 
but as the " female" prothallia bear occasionally antheridia. it 
cannot be said that a sex-determining chromosome is present. *■ e 


mean " 


J — -■— «.v -.-.-wa* *,i*_r .*..*..* WW* A A Hk. tlV %*> K, r ^^v--* ~- m 

In a foregoing paragraph reference was made to the mortaii ) 
among the spores, and to the variation in their size and apparen 
vigor. Some are larger and more healthy looking than others, an , 



abortive. It was thought at one time in this study that the 
smaller spores were the ones that produced the purely antheridial 
prothallia, but sowings were made by selecting, in so far as possible, 
the more vigorous spores (an extremely tedious process), but the 
results did not bear out the expectation. Owing to the difficulties 
of selecting individual spores, it is not maintained that the last- 
named experiment is of any great value, at least the writer does 
not lay any stress upon the results. It seems reasonable that a 
larger percentage of the smaller spores, that is, those which show 
less chlorophyll and which germinate less readily, would fail to grow 
than in the case of the larger ones, so that the possibility of the 
smaller and abortive-looking spores producing chiefly male pro- 
thallia is not excluded. However, on this point the writer can 

make no definite assertions. 


3 may be i 


of prothallia: small plants bearing only antheridia, the so-called 
male gametophytes; larger prothallia bearing only archegonia, the 
female gametophytes; and those bearing both archegonia and 


2. Archegonial prothallia, which continue growth without 

bearing a sporophyte, sometimes 


3- The gametophyte, therefore, is not strictly dioecious, and 
there is in all probability no sex-determining chromosome. 

4- It is highly probable that the development of purely male or 
female gametophytes is not dependent upon conditions of nutrition, 
but that the sexual tendency is predetermined in the spore. 
Environmental conditions, or the failure of an egg to give rise to 
a sporophyte, 


development of antheridia upon archegonial plants, which continue 



/ / — 

conditions from the dominance of the male tendency in the spore 

r the female tender 

Indiana University 

Bloomington, Indiana 




(with four figures) 

The introduction of the porous cup atmometer by Livingston has 
been followed by great activity in securing evaporation data in relation 
to plant life. Readings have usually been made with reference to a 
zero point, usually a file-scratch, on the wall of the reservoir containing 
the water. For most workers it has been impossible to visit the field 
more frequently than once a week. The" records secured were of course 
simply the total weekly evaporation. 

It is becoming increasingly clear that the record of total weekly 
evaporation can be evaluated only when we know the rate of evaporation 
at all times, and particularly when we know the maximum and minimum 
rate. This need can be met only by some form of automatic measuring 
and recording device. No such instrument has yet come into genera 

To meet the conditions of the work such an instrument must 
measure in convenient units and record in graphic form. It must 
so simple that it may be set up easily in the field and trusted to work 
with only occasional attention. Also the cost of construction must no 
be so great as to prohibit the employment of a sufficient number of the 
to secure comparative data. In attempting to meet these conditions 
as I have found them in the field, the instrument described herewit 
has been designed (fig. i). 

The device (fig. 2) consists essentially of an oscillating beam (Ah 
which may be caused to tilt by the movement of mercury in a tu e 
(B) and its connected bulbs, advantage being taken of the oscillation 
to open the stopcock (C) and to close it again. At the same time an 
electric circuit is closed which actuates the pen of the chronograph. 

In practice the whole apparatus is filled with water and the stopcoc i 
closed. Mercury is poured into the bulb b', nearly filling it, the inlet e 
is connected with the supply tank of water, and the outlet at a Wi ^ 
the atmometer. The water which evaporates from the atmometer^ 
replaced by a flow through the pipes d , c, and a from the supply in 
bulb b'. The mercury in b follows the water, partially filling b . 

Whenever the transfer of weight from b to b' has gone far enoug 
to shift the center of gravity to the right of the center of suspensio 


Botanical Gazette, vol. 50] 




2 *S 

(F), the beam A is caused to revolve until checked by the stop g'. As 
the end of A moves upward it carries with it the flange/, which strikes the 

Fig. 1 




of the stopcock and pushes it upward, thus opening the passage 
ie water supply through e, c, and a into b'. The stop g' is so 
that b does not rise to auite so high a level as b', and the mercury 




therefore flows back toward b, followed by the water which again fills 
b f . The shifting of weight toward the left causes A to return to its 




A B 

Fig. 2 

a the 

former position, the flange /' in its downward movement closing 
stopcock C. 

The point of suspension (F) is placed lower than the center of gravit} 

i 9 io] 



of the whole system consisting of A and its attached parts. In the initial 
position, as shown in the figures, the center of gravity is not only higher 
than F, but also at some distance to the left of a perpendicular through 
F. A considerable amount of mercury must therefore pass into b' 
before the center of gravity w r ill have shifted to the right of the perpen- 
dicular sufficiently to tilt the beam. The tilting of the beam carries 
the center of gravity yet farther to the right, and the beam therefore 
rests in the new position until the reverse movement of the mercury has 
carried the center of gravity to its former position in the beam. It is 
evident that if the center of gravity be raised yet higher above F, it will 
also be farther to the left when in the first position, and a larger volume 

Fig. 3 

of water will have to be displaced from b' by mercury before the beam 
will tilt. Each record, therefore, will indicate a larger amount of water. 

Advantage is taken of the relation between the center of gravity and 
the center of suspension to calibrate the instrument. A weight is placed 
upon the bar E, and by sliding it up or down the bar or exchanging it for 
another, the center of gravity is adjusted at will. The point of suspen- 
sion may also be moved. 

The atmograph, set up as shown in the photograph (fig. 4) , recorded 
the passage of 10 cc. units. By lowering the weight, the quantity could 
be reduced by half, or set at any intervening amount. Experience 
indicates that under ordinary conditions 10 cc. will be a convenient unit 
for field work with Livingston atmometers. 

Standard copper pipe and fittings are used throughout, no rubber 
being employed excepting for the flexible connection c'. Electrical con- 
nections are made from one of the binding posts shown in the photo- 
graph to the pipe e, and from the other to the flange /. The circuit is 
closed through the stopcock while / is in contact with the handle. A 
smgle dry cell is employed as the source of current. 


The atmograph may be employed in the measurement of the transpi- 
ration of an excised branch by tying the branch into a rubber tube and 
connecting with the outlet at d', or it may be used in other cases in which 
it is desired to measure the passage of water. 

For observation in the field the instrument must be inclosed in a 
box in order to preserve it from interference. If it is desirable to operate 
several atmometers at a single station, as described by Yapp, the atmo- 
graphs may be compactly arranged in a box w-ith a common reservoir 

Fig. 4 

and a single chronograph (fig. 3), as many pens being used as atmo- 
graphs. Experience has shown that the piping leading to the atmome- 
ters should be metallic. Glass and rubber are subject to many hazards, 
not only from vandals but also from the less conspicuous members of the 
fauna. Grasshoppers sometimes exhibit a very interesting, but none 
the less disastrous, adaptation of appetite to the supply of rubber tubing. 
For a series of simultaneous records a chronograph drum rotating 
in seven days will be found desirable. The records shown in fig- 4 were 
made in the laboratory upon a chronograph rotating once in six hours 
and with a group of four Livingston atmometers. The interval require 
for the evaporation of 10 cc. by the four cups has been noted to var>» 
within twelve hours, from a maximum of 113 minutes to a minimum 
17 minutes —W. L. Eikenberry, The University of Chicago. 


(with plate vii) 
So much importance has been attached at various times to the po&itio , 
shape, and development of the sporangium in Lycopodium, that any unus 
forms are of interest. The rare species L. pithyoides Schlect and ch ^ ,? 
which is figured in a former paper, 1 presents certain variations, 
stem tip is large and blunt, with a flattened apical region. The earie ^ 
stages in the development of the sporangium agree with those of previou . 

« Stokey, A. G., The roots of Lycopodium pithyoides. Bot. Gazette 44- 


described species (fig. 6). The leaves appear in close succession on the 
flattened tip, leaving no evident internodal region. The development of 
the internode afterward is very rapid, causing a conspicuous shifting of 
the position of the sporangium (fig. 7). The rapid growth of the internode, 
combined with the unequal rate of growth of the two sides of the sporangium, 
causes the foliar structure to become axial. A change of position to this 
extent is not uncommon in other species of Lycopodium, but in L. pithyoides 
there is a continued inequality in the rate of growth, so that the sporangium 
eventually takes a position on the stem entirely distinct from the leaf. 
At maturity the sporangium is not disturbed if the leaves are pulled off. 
The development in this case is the reverse of that in most species of Selagi- 
nella, in which the sporangium arises as a cauline structure, but becomes 
foliar when mature; while in L. pithyoides it is foliar in origin, but cauline 
at maturity. 

The sporangia of L. pithyoides are very large, attaining a breadth of 
2-2.5 mm., and resemble those of L. dichotomum Jacq., according to 
Bower's description and comparison, 2 although Miss Sykes's drawing,* 
while suggesting a tendency in the sporangium to be cauline, does not indi- 
cate a very close resemblance in other points. The stalk is short and rela- 
tively slender, ranging from 12 to 18 cells in diameter in sections cut radially, 
and about twice that in the other diameter. The cells are elongated and 
thin-walled, with no corner thickening and no trace of lignification such 
as Miss Sykes has described in L. clavatiim and other species. The 
vascular strand of the leaf shows no tendency to approach the sporangium. 
The leaf trace arises from the vascular cylinder 5-6 mm. below the spo- 
rangium, turning sharply upward and then making an outward bend as it 
approaches the leaf. 

The sporangia resemble those of L. dichotomum in the number of wall 
layers also. In the upper part of the sporangium there are usually four 
layers, but there may be even more owing to the irregularity in the arrange- 
ment of the middle layers (fig. 4). At the base of the sporangium the 
wall may consist of 6-8 layers. In a young sporangium (fig. 8) the order 
of division and the relation of the layers is very plainly seen, but in the 
older sporangia this regularity of arrangement is lost. The inner layer or 
tapetum is well defined as a dense, darkly staining layer. The tapetal 
cells do not show any tendency to become rounded on the inner face. In 

2 Bower, F. O., Studies in the morphology of spore-producing members; Equi- 
setineaeand Lycopodineae. Phil. Trans. Roy. Soc. London B 185:516. 1894. 

3 Sykes, If. G., Notes on the morphology of the sporangium-bearing organs of 
the Lycopodiaceae. Xew Phytologist 7:41-60. 1908. 


some almost mature sporangia the tapetum is alive, while in others very 
little older the tapetum is entirely empty, but granules and droplets are 
abundant along its inner face. The emptying process is apparently a 
rapid one, as a considerable number of sporangia were found in both con- 
ditions but none in any intermediate stages. The cells of the tapetum 
retain their shape for a considerable time and do not become crushed 
and flattened until much later. The subarchesporial pod is well developed 
(fig. 2) and slightly irregular in outline, but does not form processes in the 

The leaves afford little protection to the sporangia except in the very 
early stages, owing to the fact that they are relatively narrow (figs. 3> 5) 
and soon become recurved. The line of dehiscence, which is marked by 
smaller cells than the rest of the wall, is median (figs. 1, 3). — Alma (j. 
Stokey, ML Holyoke College, South Hadley, Mass. 


Fig. 1. — Longitudinal section of sporangium. X28. 

Fig. 2. — Tangential section of a sporangium. X26. * 

Fig. 3.— Transverse section through base of sporangium and sporangium 
stalk. X28. 

Fig. 4. — Section of part of the wall of sporangium. X345- 

Fig. 5. — Transverse section of sporangium. X28. 

Fig. 6. — Longitudinal section through leaf, showing one of the initial cells 01 
the sporangium. X725. 

Fig. 7.— Longitudinal section through stem tip, showing three stages in 
development of sporangia. X36. 

Fig. 8.— Longitudinal section of young sporangium, showing origin o« 

layers X345. 

The Third International Botanical Congress was held at Brussels 
Belgium, May 14-22, 1910. Saturday, the 14th, was the day for registra- 
tion. On Sunday, the 15th, the members of the congress assisted at a 
session of the Royal Botanical Society of Belgium held in the " dorne 
of the large building connected with the Jardin Botanique, at which se\er 
interesting papers were presented by the members of the society. Tw0 
the general sessions of the congress were held in the same room, the opening 
session on Monday morning the 16th, and the closing one on Sunday e 
22d. No regular sessions of the congress were held in the evening*, 
during the week several interesting papers on phytogeographical subj 
economic botany, etc., were given in the evening. 








sroKn 'on a copodii m 


Of the three sections into which the congress was divided for the special 
work of the week, perhaps the most important was the " Section on nomen- 
clature." The meetings of this section were held on the Exposition grounds 
in Festival Hall. 

As is well known, the Vienna Congress in 1905 selected Linnaeus 7 
Species plantar um (1753) as the starting point for the nomenclature of 
the seed plants (Spermatophytes) and vascular cryptogams (Pteridophytes). 
It also established the general principles and codified the rules which form 
the rules 0} nomenclature for plants. In dealing with the "cellular crypto- 
gams" certain problems were presented which the Vienna Congress decided 
should have special consideration, namely the question of different, later 
starting points for the nomenclature of different groups of the "cellular 
cryptogams/' and the problems connected with the nomenclature of the 
fungi possessing a pleomorphic life cycle. 

As to the starting point for the nomenclature of plants, it is well known 
that there were two opinions, as follows: (1) that there should be a single 

for the hecnnnincr of the nomenclature of all nlants: this 



starting point; (2) that there might be several different (multiple) dates 
or starting points for the nomenclature of different groups; this opinion 

'fortuity in the selection of 



Therefore the Vienna Congress wisely decided to refer the consideration of 
the nomenclature of the "cellular cryptogams" to the Brussels Congress 
in iqio, in order that these problems might be studied in the meantime. 

Since it will be several months before the complete proceedings of the 
Brussels Congress can be published, we present here, for the benefit of 
American botanists, a brief statement of the most important legislation 
enacted by the "Section on nomenclature." 

On Tuesday, the section began the consideration of the motions relating 
to the nomenclature of the "cellular cryptogams," and, with the exception 
of Thursday, which was devoted wholly to excursions, the work, together 
With the motions relating to paleobotany and phytogeography, was con- 

throughout the week. The different groups were taken up in the 
order in which they were presented in the preliminary publication, including 
the various motions, the result of the preliminary voting by the special 
ommissions, the comments of the rapporteur general, and the provisional 
''raft of rules. In general session of the section it was voted to postpone 
the consideration of the bacteria, diatoms, and flagellates, and to take 


222 BOTAMCAL GAZETTE [September 

Linnaeus' Species plantarum (1753) as the starting point for the nomen- 
clature of the Myxomycetes. 

At the opening of the session after the noon recess it was suggested that 
an adjournment of the session be taken for an hour in order to allow all 
the specialists in the different groups of cryptogams who were present to 
hold an informal conference for the purpose of agreeing upon recommenda- 
tions as to the dates for the starting points of nomenclature which would 
be acceptable to them. These recommendations were presented to the 
section on Tuesday afternoon and Wednesday morning, and were adopted 
as rules without further discussion, no one expressing a desire to discuss 
them in general session. The majorities 4 in favor of the different votes 
were very large, in the case of fungi, for instance, 130 to 4. The dates 
adopted by the Brussels Congress, therefore, for the starting points for 
the nomenclature of the " cellular cryptogams' 7 are as follows: 

Myxomycetes.— Linnaeus, Spec, plant.] 1753. 

Fungi.— Fries, Syst. Myc, 1821-1832, except for the Uredinales, 



Lichens.— Linnaeus, Spec, plant., 1753. 

.,.,._^, YfW . from., */0J. 

Algae.— Gomoxt, A T ostocaceae ho mocysteae, 1892-1893; BoRNETet* l. - 
hault, Nostocaceae heterocysteae, 1886-1888; Ralfs, British DesmMto**' 
1848; Hirn, Oedogoniaceae, 1900; Linneaus' Spec, plant, i753> for ^ 
other algae except the Chroococcaceae. 

Bryophytes.— Hedwig, Spec, muse, 1801-1830, for the Mosses; Lin- 
naeus, Spec, plant., 1753, for the Liverworts. 



M^u^xy^c, in v\uh,h tut* iuiiutuneiud.1 opecies musivf nm> *** — . 

adopted for m< sses; while for liverworts, on which there exists no wor 

1 ,.1. * 



Action upon the Chroococcaceae, bacteria, diatoms, and flagellat 
was postponed for future discus -ion, partly because it was difficult to se e^ 
satisfactory works as a basis of nomenclature, and partly because 



to take into consideration zoological as well as botanical treatise 

With regard to the nomenclature of the imperfect fungi, the following 

rule was adopted: 


of th 
■t was 

therefore greater than the actual number of delegates present. On certain g 
questions the voting proceeded simply by a show of hands. 


Fungi with pleomorphic life cycle. — 1. The different successive 
stages of the fungi with pleomorphic life cycle (anamorphoses, status) 
can bear only a single generic and specific name (binome); that is to say, 
the oldest, from the starting point of the nomenclature of the fungi, which 
has been applied to the perfect stage, provided that in other respects it 
conforms to the rules. 

2. For the purpose of nomenclature it is agreed that the perfect stage 
of fungi with pleomorphic life cycle is that which bears the ascus in the 
Ascomycetes, the basidium in the Basidiomycetes, the teleutospore in the 
Uredinales, and the spore in the Ustilaginales. 

3. Generic or specific names applied to imperfect stages may not be 
used to replace a name applied to one or more species, any one of which 
contains the perfect stage. 

Citations of pre-Friesian or of pre-Persoonian names follow the rule. 
Examples: Boletus edulis Fries, not B. edit! is Bull.; Poly poms ovinus 
Fries, not P. ovinus (Schaeff.) Fries. Writers who prefer, however, may \ 'f^jy^iriAjUr( 

write Boletus edulis Fries ex Bull.; Poly poms ovinus Fries ex. Schaeff., j gj^t^Q^-^x 


It is 


among the rusts that authors who prefer to employ double names take them . 
from the names of the host plants. A recommendation which was offered 
as applying to the fungi, namely that when a new genus is published, if 
there are more than one species the author should cite one as the type 
species, or if only one species, that one to be regarded as the type of the 
genus, was adopted and made to apply to all plants. 

The expression of opinion on the desirability of having extensive lists 
of genera conservanda was so strong that it was practically unanimous, and 
commissions were appointed to prepare such lists in the fungi, lichens, 
algae, mosses, and liverworts. 

The following action was taken in regard to genera conservanda in 
the Pteridophytes, and additional ones in the Spermatophytes presented 
for consideration at Brussels. Selaginella was placed among the genera 
conservanda, while the remaining genera in the proposed list of Pterido- 
phytes were rejected. 

A commission, which had been appointed for the purpose in advance 
of the congress, carefully considered the additional genera conservanda 
among the Spermatophytes in the list proposed by Janchen, and recom- 
mended that 21 or 22 names be stricken out. The list as amended, with 
the addition of the name Wehvitschia, was adopted. 

The motion to amend the Vienna rules by striking out the clause requir- 


ing a Latin diagnosis of new genera and species was voted down Monday 
afternoon, along with several other motions of a general nature. The 
question was discussed, however, at a later time when considering a motion 
by the paleobotanists, to the effect that a diagnosis be required only in one 
of the following languages: French, English, German, or Italian. This 
discussion broadened into a general one, and although it was defeated the 
discussion showed that there was a strong sentiment against the Latin 
requirement, especially on the part of the American botanists, and the 
subject will probably be brought up again for discussion at the next 

With reference to the question of nomenclature in phytogeograph}', 
the following principles were adopted, and the commission, considerably 
enlarged, was continued. 

i. Nomenclature is to be avoided, and the expression terminology 
is rather to be employed. 

2. When technical words are employed, a clear definition of them should 
be given in the sense in which the writer uses them, and also where a term 
is used in a sense different from that in which it has formerly been employed. 

3. It is recommended to use terms taken from living (vulgar) language* 
to designate associations, etc., and reserve expressions of Greek or Latin 
origin for higher units, where there are rarely equivalents in the living 
tongues (examples: mesophytic, hydrophytic, etc.). 

4. The principle of priority has no legal value in phytogeograph} 
Terminology is very different from nomenclature, and must be subject 
change in order to bring it in harmony with the change of ideas in the inter- 
pretation of facts. 



maps was adopted. 

plants and plant associations in their relati 
ing medium or environmental conditions). 



the section recommends the use of " formation" in a wider ecological sense. 
and " association" in a more restricted, floristic sense. For exampt 
meadow, prairie, etc., are formations; but an alpine meadow on gram 
soil in central Switzerland is an association. 

8. It was decided to publish a dictionary of phytogeograph u termi- 
nology, containing all the pertinent expressions used in phytogeograp 
and floristic works, with the original definitions and bibliographic' re e 
ences, and their equivalents in English, French, and German. 




9. It was proposed that when such words as zone and region are used 
in different senses in different countries, to employ new and clear expressions; 
for example, etage (level or floor) —Hbherenregion; Tiejerenregion of the 
Germans =zone altihre or zone abysssale of the French. 

10. A commission was named for the above purpose, consisting of the 
present members, with many additions, giving the committee power to 
add still others. 

In the publication of the proceedings of the congress, the rules and 
recommendations adopted at Brussels will be incorporated in their proper 
places with those adopted at Vienna, the latter being reprinted, so that the 
rules of nomenclature for plants will be presented in a single and conven- 
ient brochure. 

The members of the commission on the nomenclature of the cryptogams 
are greatly indebted to Dr. Briquet for summarizing in a comprehensive 
manner the manv different and often conflicting views expressed bv the 

commission in their preliminary 
ered a most important service in h 


made by different speakers in French, English, and German, so that they 
were understood by all those present. 


Mangin, and Professor Engler, who presided over the different sessions, 
performed their duties in such a way as to deserve the hearty thanks of 
all the members, combining a courteous and affable manner with a strictly 
business management.— W. G. Farlow and Geo. F. Atkinson, Paris, 

May 28, iqio. 



Heredity as an exact science 

Problems of evolution have taken a dominant place in biology ever since 
the appearance of Darwin's Origin of species. During the latter part of the 
last century there was little important progress made, though there was an 
abundance of academic discussion and repeated analysis of data recordei by 
Darwin and a few other observers; but already in the closing years of the 
century a reaction had set in. With the beginning of the present century this 
reaction reached its first important expression with the appearance of UE 



study of the various factors of evolution, an epoch not inappropriately cal 
the "era of experimental evolution." Other branches of biology have exhibited 
at the same time a corresponding development. Instead of pure speculations, 
and of generalizations from observations uncontrolled by experimental con 
tions, there has been a swiftly growing demand for greater exactness in observa- 
tions and for experimentation. . , 
As an essential part of the reaction, the development of mathematica 


key to the riddles of evolution. This movement, led by a pure mathematician, 
developed a series of beautiful methods for the mathematical analysis of 
and the comparison of variations. These methods are of the greatest im P°r 
tance when rightly used, but owing to the almost invariable lack of an equa > 
keen biological analysis, the applications of these methods have led to a larg< ^ 
spurious product, whose showing of depth and accuracy has been illusive, 
because the methods are faulty, but because of their application to ca^ 
which did not supply the one fundamental assumption of homogeneity UP 
which the whole biometrical system is based. -^ 

Parallel with this movement toward the use of mathematically pr e ^ 
methods, there has been a rapidly increasing utilization of pedigree a ^f^ > 
genetic methods involved in efforts to distinguish between mutations an 


luauuna, aim in menaenan investigations in heredity, wmui ui*«— ; - o 
chiefly upon biological analysis. The antagonism of active workers wit ^ 
metric and genetic methods in the study of heredity scarcely permitte^ ^ 
be hoped that a work might soon appear which would give a sound an ^ 
balanced treatment of heredity, utilizing the results gained by both too _^ 
and genetics. On the one hand were the mathematical writings of r0 ^ ^ 
I'earson and a number of his students, in which mathematical metho 



applied to data without a preliminary biological analysis to determine whether 
they were indeed homogeneous, as they were assumed to be; on the other hand, 
the writings of Professor Bateson and many others dealt purely with alter- 
native types so different from each other that they could be classified by ordi- 
nary inspection, and ignored entirely the fundamental value, and indeed the 
necessity, of biometrical methods in the study of less divergent types. 

It has remained for Dr. Johannsen 1 of Copenhagen, himself an ardent 
student of quantitative variation by biometrical methods, harmoniously to 
combine the mathematical analysis with a biological analysis of his material, 
that enables him to present a thoroughly well-balanced treatment of heredity 
and variation in the light of all these recent refinements of method. His book 
is presented in the form of twenty-five lectures, which give the clearest and 
simplest discussion of the statistical methods and their exact significance 
which has yet appeared. His full, elementary, and illuminating explanations 
of all the biometrical methods, with simple examples showing their usefulness 
and their limitations, are a noteworthy contribution, since they render an other- 
wise difficult and to many persons distasteful subject easily intelligible to any- 
one who will seriously undertake its mastery. While explaining the meaning 
of all the biometrical methods and their importance in the exact investigation 
of variation and heredity, the author continually lays stress upon the fact that 
these methods are instruments whose usefulness depends upon the quality 
of the data to which they are applied. Much attention is given to irregular, 
one-sided, or bimodal variation curves, and it is shown that these different 
types of curves may be produced by any one of several different causes, and that 
therefore the occurrence of such curves gives no indication of the interpreta- 
tion which is to be placed upon them. The interpretation must always come 
by biological analysis. 

The biological conception which forms the keynote of the author's entire 
discussion is the permanence of the elementary types which collectively make up 
all the systematic species of plants and animals. This is essentially the same 
conception as the " elementary species" of DeVries, but by Johannsen it 
is given an experimental support of such consistency and magnitude as to place 
it upon a much firmer basis than it has had heretofore. The existence of such 
permanent types was first given a biometrical demonstration by JOHAKNSEN 
in his Ueber Erblichkeit in Popidationen und in reinen Linien (1903) , and the 
present work is in large measure an elaboration and application of the conclu- 
sions reached in that paper, supported by the results of a number of added 
generations and additional critical experiments. 

The author's keen analysis of the different kinds of types is having a large 
influence upon present discussions of heredity and evolution. For the elemen- 

1 Johaxxsex, \V., Elemente der exacten Erblichkeitslehre. Deutschen wesent- 
hch erweiterte Ausgabe in funfundzwanzig Vorlesungen. pp. vi-f 516. figs. 3*- J ena: 
(iustav Fischer. 1909. 


tary forms — whether "elementary species" or "varieties" in the DeVriesian 
sense — he has proposed the convenient term biotypes. He also distinguishes 
them as genotypes, that is, collections of individuals having like germinal 
characters to distinguish them from mixtures of individuals having like external 
characters, but of unlike germinal composition. To these latter mixtures, 
whose uniformity is only apparent, he gives the name phenotypes. 

Heredity is recognized as only one of many factors involved in evolution, 
and the author does not even attempt to enumerate the other factors, discuss- 
ing briefly only such other evolutionary factors as are directly related to the 
question of pjermanence of types, for example, natural selection, Weismannism, 
direct influence of environment, inheritance of acquired characters, etc. He 
dismisses all which are not based upon a clear distinction between the different 
kinds of variation and the different kinds of types as purely speculative, and 
therefore as having no place in the exact science of heredity. 

The author recognizes that there may be several ways in which new biotypes 
come into existence, for example, by changes in the characters of the genes 
or determiners, loss of genes, combinations of different genes through hybridi- 
zation, etc., but devotes relatively little attention to this phase of the subject. 

While less than ioo pages are given to the results of genetic studies in 
mutation and hybridization, these subjects are treated in a very satisfactory 
manner. It is impossible in so short a space nearly to cover these fields in 
detail, and the method used is the same as in the biometrical part of the book, 
appropriate examples being introduced to illustrate each phase of the subjec , 
no attempt being made to give a comprehensive account of the many investiga 
tions which have been made. With a detailed account of the recent wor 
Mendelian heredity, we are fortunately supplied at the present time by Bat - 
son's most recent book 2 on the subject, which, together with this book y 
Johannsen, gives us a practically complete discussion of what has been 
in the study of heredity by exact scientific methods. 

While Johannsen's book is written in German, and is therefore less acces- 
sible to the entire English-reading public than might be wished, he has f° rtunat ^ J 
written with great simplicity and directness, and thus his work is more eas^ 
read than is often the case. The author has been criticized by German revie *^ 
because of certain mistakes in German idiom, but the slight departures W 
the pure German idiom always occur in the direction of the Danish I *°^ 
which is more nearly like the English; consequently, his book is more ea^ 
read by English readers than it would be if it followed the German idiom m 
closely. The value of the work is greatly enhanced by the fact that the aut ^ 
discussions are illustrated at every point by detailed examples from nib o 
incisive experiments. It is, therefore, a treasure-trove of new **° W ^ 
leading to conclusions which are certain to have a large place in the 

2 Bateson, W., Mendel's principles of heredity, pp. xiv+39^ P ls ' 6 ' ***' 
Cambridge: University Press. 1909. 



discussions of heredity and evolution. The book is worthy of a place beside 
Darwin's Origin of species and DeVries's Mutationstheorie, and is certain 
to be rated as a classic example of the new spirit which has entered into bio- 
logical investigation in the beginning of the twentieth century. — Geo. H. 

Outlines of bacteriology 

The scope of a work on bacteriology by Ellis 3 is outlined in the following 
sentence of the Introduction: "This book is intended to serve as an introduction 
to bacteriology in all its branches, though more attention has been bestowed on 
that aspect of the subject which is of the most interest to students of technical 
and agricultural bacteriology." 

Bearing the above introductory sentence in mind, the reviewer is the more 
impressed by the apparent "errors of omission and commission," the lack of 
facile expression, misstatements of fact, and in places the absence of a knowledge 
that denotes real intimacy with certain phases of the subject to be presented. 
And it is marvelous that the firm of Longmans, Green & Co., should have under- 
taken the publication of the volume. 

Without touching upon numerous smaller mistakes and errors, attention 
may be drawn to the prominence given the author's views upon the supposed 
very general flagellation of the members of the Coccaceae (pp. 19, 20). It might 
be well to point out that those views are not yet by any means concurred in by 
many eminent authorities. 

The crudity of the method recommended for the observation of the germina- 
tion of spores (p. 32) is brought into relief when compared with the superior 
advantages afforded by the use of the hanging-drop, or the hanging agar-block 
on the warm stage. 

In the light of the best American practice, the method advised for the handling 
of gelatin petri dish cultures is incomparably cumbersome and unnecessary 
(P- 47); while the strictures passed upon the use of agar under similar conditions 
are scarcely warrantable (p. 49). 

In his discussion of the effect of the electric current upon bacterial life, it 
would seem that the author had not read far enough afield, else the statements 
made by Professor J. Behrens in Lafar's Handbuch d. techn. Mykologie (vol. 
I, sec. 99, p. 455) would have vastly modified his opinion. 

Page 61 gives undue prominence to Meltzer's ideas regarding the adverse 
action of moderate degrees of continuous vibration upon bacterial cells. It may 
be said that those views no longer carry weight. 

In the chapter on sterilization (p. 85) are to be found the following remarkable 
statements in reference to the sterilization of the air of a room: "It may at once 
be stated that no gas is a disinfectant Obviously, therefore, if we wish 

* Ellis, David, Outlines of bacteriology (technical and agricultural). 8vo. 
PP- xn+262. figs. 134. London: Longmans, Green & Co. 1909. 7s. 6d. 


to purify a room, our only chance consists in spraying the walls, the floors, and 
the objects in the room with a reliable liquid disinfectant. This tends to purify 
the places from whence the air derives its supply of bacteria." The admirable 
work of Chick on the action of disinfectants receives no comment, whereas 
Miquel's work, so long out of date, is given great prominence. 

In the chapter on sewage and sewage disposal (p. 239) are to be found remarks 
that in this country at least would be deemed erroneous, improper, and inadvisable. 
For instance, who would willingly subscribe to the following: "There is great 
similarity between Bac. typhosus and Bac. coli communis, an organism which is 
very common in sewage, and which is strongly suspected of being the cause ot 
epidemic diarrhea, though positive proof is still wanting" ? Dr. Ellis condemns 
the methods of killing off bacteria in sewage by antiseptics as "not practicable, 
whereas Rideal in his own country, and Phelps and Carpenter and others 
in the United States, have shown how valuable an agent is calcium hypochlorite 
in this direction. 

In the section heading "Disposal without purification" (p. 247), discussing 
the disposal of the contents of cesspools, the author writes: "The other [insoluble] 
substances that in larger places usually find their way to the sewage drain are 
thrown broadcast on to any convenient spot, such as a roadside or a neighboring 
common. This method is efficient enough for very small places, though it mus 
detract somewhat from the healthiness of village life." It is to be regretted that 
Dr. Ellis puts himself on record as condoning such a practice; no matter un 
what conditions of life, the practice is sufficiently vile and unsanitary to be mos 
strongly condemned. 

It is very noticeable that no chapter on the biological methods of water pun 
tion found a place in the book, despite its importance to technical students. 

One cannot turn over the pages of the book without remarking upon the int 
cusable crudeness of some of the drawings, which actually mar what other" 
makes a most presentable volume. — Norman MacL. Harris. 

Vegetable proteins 


Another of the monographs on biochemistry edited by PLiMMERand HoPKl * 
just appeared,* and deals with vegetable Droteins. It is hardly necessan 


k mor 

authoritatively upon this topic than Osborne. Plant physiologists, of « ° 
fundamental knowledge in several accessory sciences is demanded, are su 
welcome a work of this kind. It shows directness and force that comes from 
author, who is the greatest producer in the subject he is discussing. T e 
ography consists of 608 citations. jl . 

A list of the chapter headings will give a good idea of the scope ot m 
historical review; occurrence of proteins in different parts of plants an 

4 Osborne, Thomas B., The vegetable proteins. 8vo. pp. xiii -K 12"- 




general characteristics; isolation and preparation of seed proteins; basic and 
acid properties of proteins; solubility of vegetable proteins; precipitation of 
vegetable proteins; denaturing of vegetable proteins; physical constants of vege- 
table proteins; products of hydrolysis of vegetable proteins; classification of 
vegetable proteins; some physiological relations of vegetable proteins to the ani- 
mal organsim and the biological relations of seed proteins to one another. 

The plant physiologist will welcome this work especially, for most discussions 
of proteins deal in the main with animal proteins. — William Crocker. 


A new catalogue of Connecticut plants. — The Connecticut Botanical Society, 
through a committee of six of its members, has issued recently a Catalogue 0} the 
flowering plants and ferns of Connecticut. 5 The publication has been modestly 
termed a catalogue, but it is far more than a mere list of plants of the state. The 
scientific and common names of the plant are given, as well as limited synonomy, 
habitat, distribution, and citation of exsiccatae; and often a note on the economic 
import of the species is added. 

In the sequence of families and in nomenclature the work accords with the 
seventh edition of Gray's Manual. A statistical summary gives the following 

composition of the flora: number of families 134, genera 621, species 1942, varieties 
and forms 286; a total of 2228 recognizably distinct plants; and approximately 
four-fifths of these are indigenous to the state. 

The work is an important one to the taxonomic student; and the collaborators 
have done a commendable service to their state in recording, in available and use- 
ful form and with a high degree of accuracy and completeness, their intimate 
knowledge of the Connecticut flora.— J. M. Greenman. 

North American Flora. 6 — Volume XXV, part II, is devoted to the Geraniales, 
as follows: Tropaeolaceae by G. V. Nash, Balsaminaceae and Limnanthaceae 
by P. A. Rydberg, Koeberliniaceae by J. H. Barnhart, Zygophyllaceae by 
A. M. Vail and P. A. Rydberg, and the Malpighiaceae by J. K. Small. Four 
new genera of the Malpighiaceae are proposed, namely, Adenoporces, Callaeum, 
Rosanthus, and Banisteriopsis. Several new species are described; these are 
distributed among the following genera: Impatiens (3), Fagonia (3), Guaiacum 
(3), Kallstroemia (6), Mascagnia (2), Hiraea (1), Triopteris (1), Tetrapteris 
(2), Banisteriopsis (2), Banisteria (2), Stigmaphyllon (1), Thryallis (1), and 
Ualpighia (5).— J. M. Greenman. 

5 Graves, C. B., Eames, E. H., Bissell, C. H., Andrews, L., Harger, E. B., 

and Weatherby, C. A., Committee ox the Connecticut Botanical Society. Catalogue 
of the flowering plants and ferns of Connecticut growing without cultivation. Bulletin 

No. 14, State Geological and Natural History Survey. 8vo. pp. 569. Hartford, Conn. 

6 North American Flora, vol. XXV, part II, pp. 89-171. New York Botanical 
Garden. 19 10 . 



Current taxonomic literature. — L. Abrams 


1910) has described 4 new species of flowering plants from California. — O. Ames 

_ a j* tt A 

(Phil. Journ. Sci. Bot.' 4: 663-676. 1909) in continuation of his sti 
pine orchids records 30 species, n of which are new to science. 
(Fern Bull. 18:9, 10. 1910) records a new variety of Lycopodium tristachyum 
Pursh from New Hampshire. — I. Boldingh (Recueil Trav. Bot. Need. 6:1-36. 
1909) under the title "A contribution to the knowledge of the flora of Anguilla 

I.)" lists about iqo species of flowering plants. The list is based on col- 



F. B0R- 

gesen (Bot. Tidssk. 30:1-19. pis. 1, 2. 19 10) has published the results of his 
studies in the Florideae collected in the waters of the Danish West Indies, describ- 
ing and illustrating 5 new species. — M. Burret (Bot. Jahrb. 44:198-238. 1910) 
presents the results of an extended taxonomic study of the genus Grewia, accom- 
panying the same by a key to the African species. — B. Chabaud (Rev. Hort. 
Paris 82:58-60. fig. 18. 19 10) has published a new species of palm (Sabal 
uresana) indigenous in the state of Sonora, Mexico.— H. Christ (Rep. Nov. Sp. 
8:17-20. 1910) has published 7 new species of ferns from Costa Rica.— A. 
Cogniaux, A. Lingelsheim, F. Pax, and H. Winkler (ibid, 1-6) under the 
title "Plantae novae bolivianae IV" have published 11 new species and 2 new 
varieties of Bolivian flowering plants. — A. Engler (Bot. Jahrb. 44- I 37 -I 55- 
1910) has published 31 new species of the Burseraceae from Africa; the same 
author (ibid. Beibl. 101, p. 34) proposes a new genus (Scirpodmdron) of the 
Cyperaceae from Africa.— E. Gilg (ibid. 15-19) describes and illustrates a new 
species of Marckea (M. Pecholtiorum) from Brazil.— W. Fawcett and A. B. 
Rendle (Journ. Bot. 48:106-108. 1910) in continuation of their studies on the 
orchids of Jamaica have published 4 new species— E. L. Greene (Leafl. Bot. 
Obs. and Crit. 2 : 25-88. 1910) has characterized about 90 new species of flowering 
plants mostly from western United States.— E. Hassler (Bull. Soc. Bot. Geneve 



raguay. The 

led several new species and varieties of flowering plants from South America. 
Two new genera of the Malvaceae are proposed, namely Blanchetiastrum from 
Brazil, and Bastardiopsis from Brazil and Paraguay.— A. von Hayek (Oeste- 
Bot. Zeitschr. 60:89-93. 19 io) has characterized a new genus (Degenia) of e 
Cruciferae, based on Lesquerella velebilica Degen— A. A. Heller (Muhlenberg^ 
6:13-32. 1910) records the results of further studies on the "Nevada lu P ,n ^ t 
and includes descriptions of 4 new species.— R. H. Howe, Jr. (Bull. Torr. o. 
Club 37: 1-18. pis. 1-7. 1910) gives a revision of the genus Usnea, as re P^ se ^. E 
in North America north of the 15th parallel, recognizing 4 species.— E. R° E '^ 
(Rep. Nov. Sp. 8:16, 17, 165-167, 196-199. 1910) has published 6 new specK 
and 4 new varieties of Cuphea from the Lesser Antilles and Paraguay. 



Kranzlin (ibid. 97, 98) has described 2 new species of orchids from the Philippine 
Islands.— G. Kukenthal (ibid, 7, 8) under the heading "Cyperaceae novae I" 
describes new species and varieties in the genus Carex, some of which are from 
America.— K. Krause (Bot. Jahrb. 44: Beibl. 10 1, pp. 9-14. 1910) has published 
several new species of Araceae from South America and the Philippine Islands — 

Th. Loesener (ibid. 44: 156-197) gives a synoptical revision of the genus Solaria, 
recognizing 64 species, of which 25 are new to science. — W. Moeser (ibid. 239- 



H. E. 

and several varieties new to science,— R. Pilger (ibid. Beibl. 10 1, p. 7) has 
published a new species of Valeriana from southern Brazil. — J. W. Palibine 
(Bull. Soc. Bot. Geneve II. 2:17-21. 1910) presents an article on the subsection 
Baicalia of the genus Oxytropis, describing one new species and transferring 
Arogalliis Bellii Greene to Oxytropis— F. Pax (Rep. Nov. $p. 8: 161, 162. 1910) 
has published 4 new species 
Petersen (Bot. Tidssk. 29:345-429. 1909) under the title "Studier over Fersk- 
vands-Phykomyceter" describes several new species and proposes one new 
genus (Pythiomorpha) from Denmark.— A. Pflle (Recueil Trav. Bot. Neerl. 
. 6:251-293. 1909) in continuation of his-studies on the flora of Surinam publishes 
a second article including 18 new species of flowering plants. — C. B. Robinson 
(Phil. Journ. Sci. Bot. 4:687-698. 1909) gives a synoptical treatment of the 
Philippine Boraginaceae, recognizing 10 genera to which are referred 2 1 species. 

P. A. Rydberg (Bull. Torr. Bot. Club 37: 127-148. 1910) under "Studies on the 


combinations in the Compositae.— F. J. Seaver (Mycologia 2:48-92. pis. 20, 21. 
1910) continues his monographic treatment of the Hypocreales of North America, 
publishes several new species, and proposes two new genera (Chromocrea and 
Chromocreopsis)—H. and P. Sydow (Ann. Mycol. 8:36-41. 1910) have pub- 


W. Tranzsche 

(ibid. 1-35) in an article entitled "Die auf der Gattung Euphorbia auftretenden 
autocischen Uromyces-Arten " has published several new species of fungi para- 
sitic on different American euphorbias.— P. Wilson (Bull. Torr. Bot. Club 37*-85, 

86. IOTO^ HpcrriKoc o mm» o*^.™^ ^f A **.„.„,;<. f*-^«-. 


new combinations in the genus Zanthoxylitm.—E. O. Wooton (ibid. 31-4 1 ) * n 


state, 5 of which are described as new; a key to the species accompanies the 
descriptive matter.— Different authors (Kew Bull. 55-60. 19 10) under the title 
"Diagnoses Africanae XXXIV" have Dublished several nevi 


a new genus (Megabaria) of the Euphorbiaceae.— J. M. Greenman. 

Respiration. — Kuijper? has studied the effect of various temperatures on 
the amount of C0 2 produced by the seedlings of Lupinus luteus, Pisum sativum, 

* Kuijper, J., Ueber den Einfluss der Temperatur auf die Atmung der hoheren 
p flanzen. Extrait Rec. Trav. Bot. Neerland. 7: pp. 109. pis. 3. 19 10. ' 


and Triticum vulgar e, with the view of testing the applicability of the ideas 
set forth in Blackman's 8 "optima and limiting factors" to the process of respira 
tion. In general the experiments were run six hours, and the C0 2 production 
determined for each hour. Up to io° the C0 2 produced per hour was constant 
for the six hours. At higher temperatures, up to 20 , there was a rise for four 
or five hours. This rise is hard to explain in the light of the facts that the seeds 
were germinated at approximately 20 , that it occurred regardless of the age 
of the seedlings, and that it was followed by a corresponding fall. At constant 
temperatures above this, up to 40 , a fluctuation in C0 2 production was apparent. 
Especially between 30 and 40 9 this manifested itself by a rapid fall for the first 
two hours, followed by a later rise ranging over one to two hours, followed by a 
later continuous fall. The author assumes that two distinct processes are differ- 
ently affected by the continuous high temperature: one early depressed, marking 
the fall; the other later stimulated, showing the rise. This fact he suggests may be 
related to the double nature of C0 2 production in respiration noted by Palladia, 
in which he assumes the action of oxidase on the one hand and of carbonase on 
the other. At still higher temperatures there is a continual fall in C0 2 production. 
The temperature at which one type of behavior changes to another is determined 
by the nature of the stored food, as the following tables show: 

Lupinus Pisum Triticum 

Rise noticeable 15-20 20-25 3°° 

Fluctuation 20-25 30 35 

Continual falling 25 35 4°° 

Protein 37% 22% i 2 % 

Starch none 54% 74% 

The Van't Hoff law applies for Pisum and Triticum from o°-20°, and 
Lupinus up to 25 . The coefficient for a temperature difference of io° lies between 
2 and 3. The continual falling in C0 2 production with continual exposures a 
higher temperatures agrees with Blackman's results. Blackman found that in 
photosynthesis the initial rate at any given high temperature (30-40°) coU 
figured in two ways, giving agreeing results: by applying the Van't Hoff coe ^ 
cient to the measurements at lower temperatures where the rate is constan , ^ 
by taking several later determinations at the given temperatures and from t est 
extrapolating the initial value. Kuijper finds that these methods will not a«W 
to the CO a yield in respiration, because of the appearance of the two anUg ? n * 
factors at temperatures between 30 and 40 , and because of the non-apph ca ° 
of the Yax't Hoff law at the higher temperatures.— William Crocker. 

Permeability.— Czapek* has published a preliminary article upon the ^ 
of various reagents upon the precipitation of tannins in plant cdb b *f^ 
of caffein and ammonium carbonate. If slices from the epidermis of £< 


•Czapek, F., Versuchs iiber Exosmose aus Pflanzenzellen. Ber. Deutsc • 

Gesell. 28:159-169. 1910. 


leaves are placed in w/1000 acid for 24 hours, and then treated with caffein 
solution, no tannin is precipitated; while the controls kept in water show a 
heavy precipitate. The acid solution show r ed a strong tannin test with iron salts. 
Czapek concludes that the acid renders the Plasma haul permeable to tannins, 

and that they gradually diffuse out. 


critical (minimum) concentration for inducing marked permeability. This is 

the concentration at which Kahlenberg and True found growth to cease. 
Czapek believes the stoppage of growth is directly due to reduced turgor, 
resulting from the induced permeability of the protoplasm to contained sub- 
stances. He recognizes that injuries appear in concentrations below the critical 
concentration for permeability. Various other substances give similar results. 
The critical concentration for phenol is 0.58 per cent or n/16; for resorcin, 
di-hydric phenol, 2.85 per cent or w/4; for pyrogallol tri-hydric phenol 11/4. 
Among alcohols, the critical concentrations were methyl 15 per cent, ethyl 
io-ii per cent, normal and iso-propyl 4-5 per cent, iso-butyl 1-2 per cent, 
amyl o . 5 per cent. In acetic acid the critical concentration is below that deter- 
mined by its acid properties. The effect of external conditions upon the perme- 
ability of protoplasmic and other plant membranes is a subject that deserves 
much attention from plant physiologists. Animal physiologists are teaching 

us much in this field. — William Crocker. 

Morphology of Sciadopitys. 


tophytes and embryo of Sciadopitys, one of the peculiar conifers of eastern Asia. 
The microspores are binucleate at shedding, and are received upon the so-called 
pollen cushion, which is the tip of the nucellus differentiated into a loose tissue of 
large thin-walled cells. During the first season only one more division occurs, 
that of the generative cell into the body cell and a free stalk nucleus. In the next 
season the body cell passes toward the tip of the pollen tube, which has entered 
an archegonial chamber, and there produces two unequal male nuclei. The 
formation of the megaspore tetrad is peculiar. The division of the mother cell 
nucleus is not accompanied by wall formation; but the subsequent division of 
the two daughter nuclei is; so that the tetrad comprises three cells, the middle 
one of which is binucleate. In these divisions the chromosome numbers prove 
to be 8 and 16. The innermost megaspore is the functioning one, enlarging very 
much, and becoming invested by the usual zone of nutritive cells. The female 
gametophyte is developed as in the majority of conifers, and there are four or 
six archegonia, each with its own investment of jacket cells and a deep archegonial 
chamber. A ventral nucleus is formed, but no ventral canal cell. In the develop- 
ment of the embryo, the free nuclei pass to the base of the egg at the four-nucleate 
stage, and the proembryo finally consists of three tiers of cells and one tier of free 



while the embryo-forming tier becomes a mass of at least sixteen cells.— J 


LAWSON, A. Axstruther, The gametophvtes and embryo of Sciadopitys ver- 

ticdlata. Annals of Botany 24:403-421. pis. 2Q-31. 1910 



The structure of Podocarpus spinulosa. — Brooks and Stiles 11 find the 
structure of the stem and leaf of Podocarpus spinulosa similar to species studied 
by Penhallow and Worsdell. The wall of the microsporangium is described 
as similar to that of Saxegothaea and Araucaria except that the dehiscence is 
oblique. The male gametophyte agrees with P. ferruginea and P. dacrydioides, 
described by Jeffrey and Chrysler, having a prothallial complex of eight cells, 
and occasionally the appearance as of a second derivative of the generative cell. 
The course of the vascular bundles in the ovulate sporophyll is studied in detail 
and compared with other forms. 

That Podocarpus is a specialized offshoot from Saxegothaea-likt ancestors 
is confirmed by the presence of less mesarch wood than in Saxegothaea, the loss 01 
function of some of the resin canals, and the specialized ovulate structure with 
reduction in size and number of scales. The independence of the vascular supply 
of the ovule from that of the scale is explained by the greater importance of the 
ovule in Podocarpus. The authors regard Podocarpineae as a natural group, 
with no very definite connection with Taxineae, no evident relation to Abietineae 
but with a probable connection with Araucarineae. — -Mary S. Young. 

Evolution of plants. — In his presidential address of 19 10 before the Lin- 
nean Society of London, Professor Scott selected as his subject ' Some 
modern ideas on the course of evolution of plants." It is an outline of the 
present status of opinion in reference to the evolution of vascular plants, 
especially as developed by the recent rapid increase of knowledge of paleo- 
botanical material, and is in part a confession of faith. The author evidently 
believes in the homologous origin of the alternation of generations, and regar s 
the sporophyte of the pteridophyte as developed directly from the thaUophyte 
body. Special attention is given to the views of Lignier in connection wi 
this thalloid origin of the cormophyte. The classification of vascular plants 
proposed in the new edition of his Studies in fossil botany is outlined, and t e 
gymnosperm relationships are discussed; while Bennettitales-like forms are 
still put forward as representing a possible origin of the angiosperms. Perhaps 
the main thesis of the address is to illustrate among plants the theory 
Gaskell, developed in a discussion of the origin of vertebrates, that eac 
successive group has arisen from some member of the highest group existing 
at the time."— J. M. C. 

Evolution of Pinus — Bailey 12 has presented the anatomical characters 

- Thecreta- 




"piriform" lateral ray pits, absence of marginal ray tracheids, and abun a 

" Brooks, M. A., and Stiles, W., The structure of Podocarp spinulosa. Anna 
of Botany 24:305-318. pi. 21. 19 10. 



Nat. 44:284-293. 1910. 

iqio] CURRE 

2 37 

tangential pitting of autumnal tracheids. The change in the living pines is 
seen in the disappearance of thick- walled ray cells, the presence of large com- 
pound ray pits, the development of ray tracheids, and the loss of tangential 
pitting of autumnal tracheids. The type of hard pines represented by P. 
resinosa in North America and P. silvestris in Europe is "the most highly 
developed and specialized condition among living pines." The nut pines (of 
North America and Asia) have piciform lateral ray pits and thick-walled ray 
cells, and in these features they are the living pines that approach most nearly 
to the cretaceous pines. The hard pines of the United States, with the excep- 
tion of P. resinosa, show a great range of variation from piciform to compound 
lateral ray pits; and the soft pines present a parallel series of gradations. 
J. M. C. 

Non-available water. — Bovie 13 has tested the effect of salts upon the non- 
available water in a soil of crushed quartz. Aside from a full nutrient solution 
of 0.2 per cent, various amounts of NaCl, ranging from 0.05 to 0.6 per cent, 
were added. To 100 grams of soil 20 grams of solutions were added. After 
the plants had grown considerably and the soil moisture was nearly exhausted, 
the cultures were placed in a special drying chamber of relative humidity of 
o.i. Soil was tested for contained moisture when the foliage began to wilt, 
and when it showed drying. The remaining water was essentially the same 
regardless of the amount of salt present. Assuming that none of the salts 
are absorbed by the plants, Bovie finds that the soil water, at the close of the 
experiments, in some cases would contain more than 300 per cent of salts, and 
that much of it must be in the solid form in spite of greatly increased solubility 
in the thin water films. He also offers some evidence for the movement of 
water in soils of low water content in the form of vapor, a thing already empha- 
sized by various workers. — William Crocker. 

Transpiration stream.— Zijlstra 14 finds that lowering the temperature of 
sections (20 cm. in length) of stem of intact plants to o° C. for several days, 
even under the most favorable conditions for transpiration, does not lead to 
wilting of the foliage. It is assumed that this renders the living cells of the 
zone comparatively inactive, without injuring them and without producing 
injurious or blocking material. The results are contrary to those obtained by 
the same method by Urspruxg, who used the results as an argument for the 
necessity of the activity of living cells to the continuity of the transpiration 
stream. Zm 


of 0.1 per cent Saureviolett (Grubler) in living and dead stems. He also 

13 Bovie, William T., The effects of adding salts to the soil on the amount of 
non-available water. Bull. Torr. Bot. Club 37:273-292. 1910. 

14 Zijlstra, K., Contributions to the knowledge of the movement of water in 

Plants. Reprint from Koninklijke Akad. Wetenschappen te Amsterdam. 1910:574- 




studied the behavior of plants in which lateral movement of water was made 
necessary by the ordinary method of incisions on opposite sides. Only the 
first set of experiments throws any new light on the question of the rise of sap. 
William Crocker. 

Phylogeny of Filicales.— Bower 15 has begun the publication of a series of 
studies in the phylogeny of the Filicales, and the first paper presents the singular 
and poorly known genus Plagiogyria. It is grouped among the Polypodiaceae, 
and even merged with Lomaria, but the position is anomalous on account of the 
oblique annulus. Several species were secured, both oriental and occidental, and 
the results may be considered fairly representative for a genus of eleven species. 
The conclusions are that the genus is quite a distinct one, deserving to stand apart 
from Lomaria or any other genus; that it is relatively primitive, as shown by 
stelar structure, leaf traces, venation, etc.; that it has resemblances to the Simplices, 
but that its characters indicate that it is rightly placed among the Pterideae; and 




/ — j — x A 

with its primitive characters and its affinity to the Pterideae, shows that the Mixtae 
have probably come directly from the Simplices, rather than by any of the Gra- 
datae— J. M. C. 

Geotropism and split stems. — Schtscherback, 16 working in Pfeffers 
laboratory, has carried on a series of experiments on the geotropic reactions 
split stems. In the main, Lupinus alb us was used, for in this 
strains between cortex and pith are slight. Splitting cuts the rate of growth even 
in the vertical position, but two equal halves grow with equal speed in that posi- 
tion. If the two halves are placed horizontally, with the split faces together an 
horizontal, the lower half is favored in its growth and the upper half greatly mhi - 
ited, or even entirely stopped in most cases. Restoring the two halves to t eir 
normal vertical position restores them to equal growth, and rotating them I 

[y gives an interchange 

of rates of growth. . The same geotropic inhibition and favoring of growth occur 
in the two separated halves of the stem as occur in their intact condition. 

William Crocker. 

Relation of living cells to the rise of sap.— Rorhardt, 17 in his voluminous 
article on the part played by living cells in the rise of sap, has followed the met o 
of wearying the reader with the publication of his notebook, instead of preparing 
a concise statement of his methods and results. Of the 125 different species 

, monocotyl and dicotyl 

'5 Bower, F. O., Studies in the phylogeny of the Filicales. L Plagiogyria. Anna 

posit io 


of Botany 24:423-450. pis. 32, 33. figs. 5. 1910. 

wmay ^4:423-45°. f». fc 33- fig s - 5- I9"- eln 

16 Schtscherback, Johaxn, Die geotropische Reaktion in gespaltenen Stenger 

Beih. Bot. Centralbl. 251:358-386. 1910. 

»7 Rorhardt, P. A., Leber die Beteiligung lebender Zellen am Saftsteigen >' 

Pflanzen von niedrigen Wuchs. 



families that he has investigated, he finds that the killing of a short zone of a 
stem or of a petiole results very soon in cutting down the water transport to the 
parts above, and leads to wilting — the longer the killed zone, the quicker the wilt- 
ing. He claims that there is no blocking or interfering with the water path, but 
that the reduced water movement is due to the lack of living cells in the zone 
killed. This indicates that the continuous rise* of sap in small plants, as has been 
shown to be the case in tall trees, is dependent upon the action of living cells 

William Crocker. 

Seedling structure of Gnetales. — Hill and DeFraine 18 conclude their account 
of the seedling structure of gymnosperms by a presentation of Gnetales. A 
short cotyledonary tube is formed in every case; the cotyledonary traces are two 
in Ephedra, four (in two pairs) in Welwitschia, and four or five in Gnetum; and all 
the traces are collateral endarch. The so-called foot in Welwitschia and Gnetum is 
described as a parenchymatous growth at the base of the hypocotyl, in the former 
genus being spadelike and with no vascular supply, and in the latter genus being 
rodlike and with numerous well-differentiated bundles. The transition to root 
structure occurs in the lower region of the hypocotyl, in Welwitschia and Gnetum, 
immediately below the foot; and in all cases the primary root is diarch. — J. M. C. 

"Apogamy" in Pteris. — Miss Stephens and Miss Sykes have examined 
prothallia of Pteris droogmantiana furnished by Boodle. In a brief note 19 they 
announce that binucleate cells are common; that there has been no nuclear 
migration, as no surrounding cell is without its nucleus; that the two nuclei arise 
from the division of the nucleus of an ordinary cell without wall-formation; that 
the two nuclei thus produced remain separate for some time and then fuse. Just 
what this performance means is not evident, but perhaps the fuller paper will 
tell. The note is called one on "apogamy," and presumably there is reason for 
knowing that these are apogamous prothallia. — J. M. C. 

Morphology of Psilotum. — Stiles 20 has investigated the anatomical structure 
of the serial shoots of Psilotum ftaccidum, one of the two species usually regarded 
as constituting the genus. Aside from certain anatomical details that distinguish 
this species from the other (P. triquetrum), the main results are that secondary 
thickening is found in this species also; that mesarch structure occurs in the lower 
part of the aerial stems; that the sporangiophore trace terminates in the central 
tissue between the three "confluent" sporangia, as had been shown for Tmesip- 
teris; and that no evidence was obtained to decide whether the sporangiophore is 
"foliar" or is "an organ sui generis:'—}. M. C. 

18 Hill, T. G., and DeFraine, E., On the seedling structure of gymnosperms. 
IV. Gnetales. Annals of Botany 24:319-333. pis. 22, 23. 1910. 

J 9 Stephens, E. L., and Sykes, M. G., Preliminary note on apogamy in Pteris 
droogmantiana. Annals of Botany 24:487. 1910. 

20 Stiles, \V.,The structure of the aerial shoots of Psilotum flaecidum Wall. Annals 
of Botany 24:373-397. pi 25. 1910. 


Removal of cotyledons. — Jacobi 21 has studied the effect of the removal 
of various amounts of cotyledonary tissue upon the rate of growth of seedlings. 
In both light and darkness, removal of one cotyledon and even a part of the 
second early but temporarily stimulates the rate of longitudinal growth of 
the stem of Phaseolus multifiorus. In Cucurbita Pepo and conifer seedlings, 
elongation of the stem is stimulated in darkness, while in light the expanse of 
the cotyledon is most increased. In Cucurbita and the pines, the cotyledons 
of course bear little food material and function as leaves. The paper adds 
little that is new. — William Crocker. 

Morphology of Cunninghamia. — In 1908 Miyake published a preliminary 

account of the gametophytes and embryo of the monotypic Cunninghamia 
which was reviewed in this journal (46:156. 1908). Now the full paper has 
appeared, 22 with its excellent plates and photomicrographs. The review 
referred to includes a synopsis of the most important results, so that it is only 
necessary to announce the appearance of the full paper. — J. M. C. 

Insect galls. — Beutenmuller 2 ^ has published two more valuable papers on 
the American gall-producing insects and their galls. In these two papers he 
takes up 44 species, including three new ones, gives complete synonomy and 
bibliography for each, and clear, comprehensive descriptions of both the galls 
and the insects. This series of papers is of very great value to both the botanist 
and the entomologist. — Mel T. Cook. 

A new disease of apples. — Lewis 24 has recently described a new species 
Endomyces (E. malt), which he finds capable of producing a slow decay of ripe 
apples. This is the first species of this family to be reported from the United 
States, and hence many will be interested in the facts he gives regarding the mor- 
phology, spore production, relationship, and cultural behavior of this new species. 


E. Mead Wilcox. 


Polarity. — Miss Freund 25 has made a study of polarity in the twigs c 
plants. The paper offers nothing new on the subject. Her work might have 
been far more conclusive if she had been acquainted with the work "of MaCAI*- 

■William Crocker. 


21 jacobi, Helexe, Ueber den Einfluss der Verletzung von Kotyieaoncu »«. - 
Wachstum von Keimungen. Flora 101:279-289. figs. 2. 1910. 

V T«U~ J l ^ -r.i__ _._ _^__i _.._*. I ™*KrvrncrpnV in CM*' 


ninghamia sinensis. Beih. Bot. Centralbl. 27:1-25. pis. 1-5. i9 IQ - 

23 Beutenmullek, Wm, The North American species of Neuroterus and their 
galls. Amer. Mus. Nat. Hist. 28:117-136. pis. 8-13. 1910; and the North Americ 
pedes of Aylax and their galls. Idem 28:137-144. pi. 14. 1910; 

E., An Endomyces from apple. Bull. Maine Exp. Sta. i7 8: -*5 4 ' 

fi$ s - 58-7 T - i9*°- 

2 5 Freund, 1 

308. 1900. 

Flora 12*79"" 

26 MaCallum, W. B., Regeneration in plants. Bot. Gazette 4<>:97- i20 > ul ~ 

263. 1905. 


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The American Sociological Society 

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Religion and the Mores William G. Sumner 

History of the American Social Science Association F. B. Sanborn 

Ranges in the Census Methods for the Census of 1910 E. Dana Durand 

1 he Outlook of American Statistics Walter F. Wilcox 

he Social Marking System Franklin H. Giddings 

l he Psychological View of Society Charles A. Ellwood 

JJuthne of a Theory of Social Motives James M. Williams 

Th p" dy of Homeric Religion Albert C. Keller 

*ne Role of Magic James Thomson Shotwell 

nttuence of Superstition on the Evolution of Property Rights, Hutton Webster 

Notes on the Recent Census of Religious Bodies George A. Coe 

* ft e leaching of Sociology James Quayle Dealey 

sociology and the State Lester F. Ward 

he Sociological Stage in the Evolution of the Social Sciences, Albion W. Small 
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A Plan of Study Intended to Develop the Student's 

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Relation of Soil Moisture to Desert Vegetation 

Burton Edward Livingston 

Longitudinal Compression upon the Pro 
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L. H- Pennington 

The ITorth American Species of Stereocaulon 

Lincoln Ware Riddle 

Briefer Articles 

Toe Epidermal 

Characters of Frenelopsis ramosissima 

Edward Wilber Berry 

Current Literature 

The University of 


William Wesley and Son, London 

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Ghe ^Botanical <3asette 

H toontbls journal Bmbracins all Departments of botanical Science 

Kdited by John M. Coulter, with the assistance of other members of the botanical staff of the 

University of Chicago. 

Issued October J 5, 1910 



Burton Edward Livingston 



MECHANICAL TISSUE IN STEMS (with two figures). L. B. Pennington - - 257 


Ware Riddle 



The Epidermal Characters of Frenelopsis ramosissima (with two figures). 

Edward Wilder Berry 305 





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Botanical Gazette 




Burton Edward Livingston 

(with four figures) 

Introduction. — Of a number of lines of study now in progress, 
looking toward some quantitative knowledge of the relation obtain- 
ing between vegetation and environmental conditions, the determi- 
nation of the march of soil moisture at Tucson has now progressed 
sufficiently far to warrant publication. The present paper deals 
with a continuation of the data upon soil moisture published by 
the writer under the title: "The soils of the Desert Laboratory 
domain" in Spalding's recent monograph on desert ecology (i). 
The data there presented comprise observations on soil moisture 
content extending over the period from October 1907 to April 
1008. The period of observation has now been extended to March 
iooq> thus including both the spring dry season and that of the 
summer rains. To make the present presentation complete, the 
observations already published are here included. 

Relation of soil moisture to other factors. — Just as the 

evaporating power of the air and the nature of the transpiring 
organs practically determine the water requirement of plants, so 
do the soil moisture content and the nature of the root system 
determine the water supply. Aside from those cases (of toxic 
soils and soils of high osmotic pressure) where the nature and 
amount of the solutes in the soil solution exert an influence upon the 
character of the root system or upon the rate of water absorption 

1 Publication from the Botanical Laboratory of the Johns Hopkins University, 

No. 16. 



by these organs, and aside from those cases where the chemical 
content of the atmosphere or the character or intensity of illumina- 
tion exert an influence upon the structure of the transpiring organs 
or upon the general tone of the plant, the simple relation between 
soil moisture and the evaporating power of the air appears to be 
of prime importance in determining the character of the natural 
vegetation. Over an area such as that here considered, where the 
air and light conditions are approximately uniform, and where the 
chemical content of the soil appears not to vary sufficiently to 
produce variations in plant growth, the possible rate of water 
supply to plant roots may be supposed to become the limiting 
condition. This rate of supply depends primarily upon the abso- 
lute amount of soil moisture and upon the water-conducting 
power of the soil. 






tion or other water entering the soil from above or laterally, and 
by the water-retaining power of the soil. Water removal from the 
soil is a resultant of evaporation at the soil surface, downward 
movement into lower layers, and plant absorption. 

Throughout the area here studied precipitation is approxi- 
mately uniform. Surface drainage, nearly uniform within the 
extent of each soil type, is quite different (because of variations 
in slope and porosity) for the four types. The water-retaining 
powers of the four soils are also different. Underground drainage 
is very different for the different soils, as is also the depth of 
permanent subterranean water, which is practically quite absent 

from two of the soils. 



Spalding above cited. Tumamoc Hill, on which the Desert 
Laboratory stands, rises abruptly from the broad valley of the 
Santa Cruz River. The valley floor slopes very gently from the base 
of the hill to the river floodplain and to the various "washes," 
tributaries of the main river channel The four soils here considered 


may be briefly described as follows. A more complete description 
is included in the author's article above cited. 

1. The soil of Tumamoc Hill is a heavy clay, underlaid by 
practically impervious rock at a depth of but 10-50 cm., and much 
broken into small pockets and irregular masses by outcropping 
rock or large rock fragments. Fully 50 per cent of the entire 
gross volume of the soil is made up of large and small, rock masses. 
A subterranean water table does not exist here, so far as is known, 
and, from the impermeability of the underlying rock both to water 
and to plant roots, it is safe to conclude that the water supply for 
this soil is derived exclusively from precipitation. The unsifted 
soil possesses a water-holding power of 48 per cent of its dry weight. 
The usually pronounced slope of the soil surface here is partially 
offset by the numerous small catchbasins formed by rock frag- 
ments, so that rain water stands in small pools over the hill and does 
not drain away quite so rapidly as though the surface were more 



the creosote bush thereon, occupies the gentle slope which surrounds 

the hill 





, the 

The surface 

about half its volume of fine angular gravel; that near the caliche 

* i • - 1 1 




pebbles. The slope, the smoothness and impermeability of the 
hardpan, the presence of rock fragments, and the loamy nature of 



like the hill soil, is not influenced by subterranean influx of water. 
Unsifted soil from near the surface possesses a water-holding power 
of only about 20 per cent. This low retaining power is not here 
due to general coarseness of the soil, but to the large proportion 
" f angular gravel, which of course holds practically no water. 

3- The soils of the wash, a broad streamway from the Tucson 
mountains, which cuts through the general valley floor (Larrea 
sl °Pe in this vicinity) near the northern base of the hill, vary from 
gravel through sand to very loamy sand. The coarser soils are 
Practically without vegetation and are of little interest in this 


connection. The loamy sand, constituting little floodplains on 
the margins of the actual streamway, is 30 cm. or more in depth 
and is underlaid by coarser material. This is the soil here con- 
sidered. It is practically without gravel admixture and has a 
water-holding power of about 25 per cent of its dry weight. It 
is almost certain (but without direct evidence) that an under- 
ground water table is present here, at least for many months of the 
year. The soil surface is nearly level and water is received from the 
superficial run-off of the hill and Larrea slope ; however, it is well 
drained below. 

4. The soil mass of the river floodplain, now completely aban- 
doned by the stream, is of unknown depth. The surface layer is level 
and composed of clay loam with a water-holding power of about 
39 per cent of its dry weight. This is about 8-9 meters deep and 
rests on sands and gravels, permanent water being met with at a 
depth of 10-15 meters. This soil is characterized by general 
absence of surface run-off and by but slow percolation to lower 
layers. The dry soil of the deeper layers, however, removes mois- 
ture from near the surface as rapidly as it can move downward by 
capillary action. 

The plant societies.— The plant societies of the area about 
the Desert Laboratory have been thoroughly described by SpAL- 


(2) . Thornber's ecologically classified list of species for this region 

(pp. 103 

112 ot the same mono 

to the student of environmental factors. The four types of vegeta- 
tion which occupy respectively the four soil types here considere 
may be briefly characterized as follows: 

1. The Parkinsonia society of Tumamoc Hill possesses more 
forms, both woody and non-woody, than any other of the four so- 
cieties here considered. Besides this form of palo verde {Parkinsonia 
micro phylla) , the group includes the giant cactus (Cereus giganteush 
the ocotillo {Fouquieria splendens), the barrel cactus (Echi- 
nocactus Wislizeni), several opuntias of both cylindrical and flat- 



small specimens of mesquite, etc. Bigelovia HartwegU occur 
here but does not attain a« ar^t mimk^rc «r a« lare^e size as on 


river plain, where it is more characteristic. Numerous low forms 
are active throughout the year, notably Encelia farinosa, Sphaeral- 
cea pedata, etc., and a great array of summer and winter forms 
are to be found at the proper season. 

2. The Larrea society of the gravelly slope exhibits, in many 
localities, almost no woody species other than the creosote bush 
{Larrea tridentata). The bushes are uniformly and openly dis- 
tributed over a surface which is otherwise nearly bare of vegetation, 
excepting during early spring and late summer, when numerous 
low forms appear. Of the desert shrubs within its climatic range, 
Larrea seems best fitted to withstand prolonged drought both of 
air and soil. Ephedra occurs upon the slope as well as in the wash, 
but makes its best growth in the latter place. 

3. The Cercidium torreanum society of the wash is characterized 
by this form of palo verde and by mesquite (Prosopis), Condalia 
lye hides, and Acacia Greggii. The three latter forms are also 
characteristic of the Prosopis society of the plain. Ephedra 
trijurca and the creosote bush (Larrea tridentata) attain here their 
most luxuriant growth, though the latter, at least, is most char- 
acteristic of the Larrea society. Aside from these and some other 
woody plants, the vegetation of the wash consists mainly of small 
forms, which are active only during the periods of moist soil, 
either in winter or summer or at both seasons. 

4- The Prosopis society of the river plain is dominated by 
mesquite, two forms of Acacia (A. constricta and A. Greggii). and 
Condalia lycioides. Annuals of many forms are numerous during 
the moist periods. The most striking of the low perennials is 
Bigelovia Hartwegii, which scarcely occurs outside this group, 
except for scattered plants in the Parkinsonia society. 

Methods of determining and of representing soil condi- 
tions.— The method used in determining the water-holding powers 
given above was to nlace the. soil in a sheet-metal cylinder about 


cm. in diameter, with 






From the weight of the 


dry and saturated soil the water-retaining power is calculated in 

the usual way (3). 

The actual soil moisture content was determined by the weigh- 
ing and drying method, the temperature of drying being 105-no C. 
The samples were taken, by digging, at depths of 15 and 30 cm. 
for all soils but that of the Larrea slope. For the latter soil, on 
account of its shallowness, the depths were 10 and 20 cm. Samples 
for each soil type were taken from the same area of a few square 
meters, the small excavations being immediately refilled. No later 
sample was taken from within 50 cm. of the soil thus disturbed. On 
December 13 irrigation water made it necessary slightly to change 
the position of the station for the soil of the floodplain, otherwise 
the stations remained the same throughout the whole period. 
Samples were taken at intervals of about 10 days. Three samplings 
were omitted for all soils during the month of June 1908. A few 
other omissions of single samples occur. 

The specific gravities of these fours soils are approximately the 
same, there being almost no content of organic matter. Therefore 
the moisture contents are calculated on the basis of the dry weight 
of the soil and are here approximately comparable. 

The careful work of Mr. J. C. Blumer, Mr. E. E. Sherff, 
and Professor J. E. Kirkwood, who performed the operations of 
collecting, weighing, and drying the soil samples, as well as that ol 
keeping the rain record, is here gratefully acknowledged. 

The accompanying graphs (figs. 1-4) have been constructed 
for each soil separately; the lesser depth is represented by a thin 
line and the greater by a heavy line. The precipitation record 


rain r< 
\ same 


taken. For easy comparison. 

graphs of soil moisture simply 
w h of each block denoting the 
relative amount of precipitation which took place during the 
corresponding ten-day period. The figures directly above these 
blocks denote the precipitation in centimeters, that is, the relative 
length of the blocks. The data from which the moisture graphs 
are constructed are placed upon the figures directly above the base 




± o 














r- oqIo 

os w 














« V* 





P h> 



a £*«. 


_ _ — * - ' '■ ■» — 

*2 < 



I- O 

«* crv 


line of these graphs. They express percentage of dry weight. 
At the base of each figure, the dates of sampling are indicated. 
The major portion of the calculations and of the drawing of the 

graphs are the work of Grace J. Livingston. 

Discussion of graphs. — -It is to be remembered that several 
simultaneous samples at the same station would be expected to 
vary to some extent, especially in the stony soil of the hill and of the 
deeper layer of the slope. Therefore it is not surprising to note 
minor fluctuations in the graphs, which do not appear traceable to 
conditions of precipitation. 

The graphs bring out clearly the lagging of soil moisture behind 
precipitation; it requires considerable time for rain water to reach 
the upper level of sampling and a still longer time for it to reach 
the lower level. A slight precipitation may not alter the condi- 
tion of the soil at either level, the water being lost by evaporation 
before it can penetrate even to a depth of 10 or 15 cm. It also 
occurs that a rain which is only sufficient to moisten the surface 
layers, thus placing them in capillary connection with the deeper 
ones which are already moist, may actually accelerate the drying 
out of the deeper layers. Thus a slight rain is sometimes directly 
deleterious to the vegetation ; it removes the dry mulch from the sur- 




From data given in Publication 50 of the Carnegie Institution 
(pp. 66, 67) it appears that a number of plants of the hill, when 
grown in pots (in hill soil), wilted with a soil moisture content of 
6-12 per cent by wet volume. On the basis of dry weight these 
numbers become about 7 per cent and 14 per cent. Boerhavia 
(one of the summer annuals of the hill) wilted in the open soil with 
a moisture content of 6-7 per cent by wet volume, or about 7-8 
per cent on the basis of dry weight. These observations were 
taken in July and August, the period of the summer rains, when the 
evaporating power of the air was far below its magnitude of May 
and June or of October and November. With greater evaporation 
r ate, the plants should of course wilt with a correspondingly 


It is probably safe to assume, there- 




fore, that during the driest months of the year the majority of the 
plants of the hill require an amount of soil moisture of 8-10 per cent. 
With a drier soil than this it is probable that most forms without 
pronounced storage organs would succumb in a few weeks. 

No determination of wilting points is as yet available for the 
other three soils, but from the well-known fact that the " non-avail- 
able water'' in soils decreases, in general, with their water-holding 
power, it is probably safe to assume that vegetation can withstand 
a considerably drier soil in the case of the slope and wash, and a 
somewhat drier one in the case of the river plain. 

From the graphs it is evident that the period of most intense 
drought occurs just before the beginning of the summer rains. In 
general, the month of June may be taken as the month of driest 
soils, and it is also the month of greatest evaporation. There- 
fore, for plants without well-developed water storage tissues, this 
month must approximate the critical period of the year so far as 
the water relation is concerned. It will therefore be instructive to 
derive the average soil moisture content for this month in the case 
of each of the four soil types. So far as I am aware, the root systems 
of all plants of this area, which are without storage tissues and which 
are not annuals, penetrate into the soil at least to the greater 01 
the two depths here considered. We may therefore consider the 
greater depth alone. The following table presents these averages 
for the four determinations taken from June 2 to July 2, i9° 8 ' 
The water-holding capacities are also given, and the water conten 
as percentage of the latter. 





in percentage 
of dry weight 


Depth in cm. 

Hill . . 





Percentage of 
dry weight 





water holding 


16. 1 

That the vegetation of the hill is the most varied and the most 
perennially active of all the plant societies of our area, agrees we 


with the high soil moisture content (almost n per cent) and the 
high percentage of the moisture-retaining power (almost 23 per 
cent) which this soil exhibits during the driest month of the year. 
It is to be remembered in this connection that the soil of the hill 
is often considerably over 30 cm. in depth, and that, in the dry 
season, the highest moisture content is to be expected just above 
the underlying rock surface. In the summer of 1904, immediately 
preceding the advent of the July rains, a soil sample from a depth 
of 35 cm. on the hill exhibited a moisture content of 17.9 per cent 
of its dry weight. From the graph of the hill soil and from this last 
consideration may be derived convincing additional evidence in 
favor of the conclusion expressed by Spalding and by the author 
(Publication §0, Carnegie Inst., p. 12) "that sufficient moisture 
is probably at all times present in the deeper layers of these soils 



excellent soil conditions exhibited by Tumamoc Hill are emphasized 



observed moisture content at a depth of 

cent, and that it was far above 11 per cent at all times excepting 

in the months of June and November 1908. 

The soil of the Larrea slope, at a depth of 20 cm., just above 
the caliche hardpan, exhibited only 3.1 per cent of moisture as the 
June average. Had the soil been sifted, the moisture content 
would undoubtedly have been markedly greater, as is indicated 
by the high percentage of its retaining power (15.5 per cent) 
which the moisture content represents. But the amount of water 


this time. 

In this vicinity, practically the only perennially active plant 
on the slope, excepting a few scattered cacti and here and there a 
plant of Fouquieria (itself hardly perennially active), is the creosote 
bush, from which the slope is named. This plant, as has been 



J forms in the region. An inspection of the graph 
lgs out the fact that the moisture content rose 
t only in February and March, in July and August 
winter of 1909. In distinct opposition to the soil 


of the hill, this soil is undoubtedly deficient in moisture for by far 
the greater part of the year. It usually supports spring and sum- 
mer annuals for a few weeks during each of its moist periods. 

The soil of the wash shows an even more inadequate moisture 
supply for the month of June than does that of the slope. This 
soil contains practically no gravel in its superficial layers and its 
moisture content at this time (2.1 per cent by dry weight) was only 
8.4 per cent of its water-holding power. From the general aspect 
of the vegetation it would be concluded that this soil was better 
supplied with moisture than that of the slope, but the graph fails 
to explain this view. The discrepancy is apparently due mainly 
to the great depth of the soil, which acts as a much more extensive 
reservoir for water than the shallow soils of hill and slope. The 
presence of underground water is doubtless effective in case of 
those trees and shrubs the roots of which penetrate deeply. The 
continuously active forms which occur here must be considered 
as much more deeply rooted than is possible on the hill and on the 

The soil of the river plain exhibits a much more favorable mois- 
ture condition, not only in June, but throughout the year, than any 
other of our four soil types excepting that of the hill. Its average 
water content for the drought period was 6.3 per cent by dry 
weight. In terms of its moisture-holding power, this becomes 
16. 1 per cent. This is a deep soil, with unknown, but surely con- 
siderable, amounts of moisture below the limits of sampling. It 
also has constant subterranean water at depths which can probabh 
be reached by tree roots. This latter fact seems to explain the 
occurrence here of mesquite, acacia, and other deep-rooting shrubs. 
In the dry seasons (early summer and late autumn) this soil shows 
very few active plants besides those just mentioned, but in the 
rainy seasons it supports a dense growth of annuals, being apparenth 
as well suited to such forms as is the soil of the hill itself. That 
many annuals of the hill fail generally to appear on the plain, and 
conversely, is perhaps to be explained by the considerably higher 
moisture content occurring during the rainy seasons on the former 
soil. It may also be related to conditions which hinder surfao 


evaporation on the hill, such as the presence of numerous rock 
fragments on or in the soil. 

An interesting question is suggested by the fact that the creosote 
bush dominates the slope and occurs on the hill, but is almost 
entirely absent from the plain, excepting along the margin where 
it abuts against the slope. No evidence as to why this shrub should 
not thrive on the plain is as yet at hand. The most vigorous 
specimens of the area occur along the lower margins of the slope, 
which agrees with a similar behavior always observed in irrigated 
specimens, in seeming to indicate that a more adequate water supply 
produces an abnormally great growth. Spalding mentions the 
physiographical relation (which of course cannot directly affect the 
vegetation) that Larrea usually dominates gentle slopes which are 
being eroded at the present time. It would seem that there must 
be some condition in the soil of the plain which is antagonistic 
to the growth of this shrub. A study of the oxygen conditions of 
these soils may possibly throw some light on the subject. The 
moisture condition alone is inadequate to explain the facts. 

The results obtained from a study of the graphs would place the 
hill soil first, as the best suited for general plant activities ; the soil 
of the plain would occupy second place, and the two other soils 
would lie rather close together and far below either of these. This 
general arrangement agrees well with the vegetational characters 
of the four soil types if we consider that the deep-rooting forms of 
wash and plain obtain moisture from levels deeper than those here 
considered. It is also in fair agreement with the simple series of 
the water-retaining powers of the four soils. From this retaining 
power alone the soils would have been interpreted in the same 
general way. This emphasizes the water-holding power as an 

easily obtained and very important factor in studies of distribu- 

The last-named factor has been used by the writer (4) with 
some success in a search for the determining conditions of forest 
distribution in Michigan. It appears that for different upland 
habitats of the same region, where the level of underground water is 
far from the surface, and the precipitation and evaporating power 

256 BOTANICAL GAZETTE * ■ [October 

approximately uniform 


water-holding power. The ease of obtaining comparative data 
for this factor entitles it to a thorough test as a soil criterion for 
habitat studies. 

The Johns Hopkins University 

Baltimore, Md. 



1. Spalding, V. M., Distribution and movements of desert plants. Publ. 
113, Carnegie Institution of Washington. 1909. 

2. , Plant associations of the Desert Laboratory domain. Plant 

World 13:31-42, 56-66, 86-93. 1910. 

3. Livingston, B. E., The relation of desert plants to soil moisture and to 
evaporation. Publ. 50, Carnegie Institution of Washington. 1906. 

4» , The relation of soils to natural vegetation in Roscommon and Craw- 
ford counties, Michigan. Bot. Gazette 39:22-41. 1905. 




L. H. Pennington 

(with two figures) 


For more than a century observations and experiments have been 
made to determine what factors cause or influence the production 
of mechanical tissue in plants. Previous to Pfeffer's time, the 
causes were supposed to be of a mechanical nature. These expla- 
nations have been shown to be inadequate. 1 Pfeffer (26, 27) 
showed that tension, pressure, or even contact may act as a stimulus 
to cause the production of more mechanical tissue. Newcombe 
(21) suggested that plants must respond to stress in a self-regula- 
tory manner by producing mechanical tissue where it is most needed. 
He also indicated the complex notion in regard to stress, and sug- 
gested that it should be subdivided. At present we use the terms 
tension or traction, and pressure or compression, to indicate two 
opposite kinds of stress 

To determine the effect of each kind of stress, much careful 
experimentation is necessary. Hegler (12), Ball (i), Hibbard 
(14), and Bordner (2) have investigated the influence of tension 
upon growing cells, with regard to its mechanical as well as to 
its stimulatory effects. Although these investigators did not 

agree in all points, their results show that tension alone cannot 
cause a very marked increase in mechanical strength. Vochting 
( 34), Hibbard (14), and Bucher (4) have contributed a little 
concerning the effects of longitudinal compression. 

Vochting reported that he fastened weights upon the tops of 
sunflower stems, some of which had been decapitated, and upon 
Savoy cabbages, all of which had been decapitated. After leaving 
the plants thus weighted for several months, he was able to discover 

Botanical Gazette, vol. 50] 

Ursprung, Biol. Centralbl. 1906. 



no increase of mechanical tissue in the sunflower stems, and but a 
slight increase in the cabbage stems, an increase not at all propor- 
tional to the weights used. 

Hibbard suspended weights upon the stems of the sunflower, 
periwinkle, fuchsia, and coleus, and found in all except coleus a slight 
response to compression by self-regulatory increase in mechanical 
tissue. He did not believe, however, that the evidence could be 
regarded as conclusive. 

Bucher inclosed both the upper and the lower portions of stems 
of the castor oil plant in plaster casts, and then fastened these 
casts so that the stems as they elongated were subjected to a longi- 
tudinal compression. All but two of his plants became bent; 
the two which remained straight were weaker in mechanical tissue 
than normal plants. 

From the few data which have been recorded, it is very plain 
that we do not have sufficient evidence to show whether or not 
compression or increased weight may act as a stimulus to cause the 
production of a greater amount of mechanical tissue in plant stems. 
General conclusions have been drawn from mere observations of 
phenomena, which may be due to any one of several influences 
or to the combination of two or more influences. The few experi- 
mental data have led to indefinite or contradictory conclusions. 
This investigation was undertaken to secure exact experimental 
data upon the effect of weight tending to produce a longitudinal 
compression in vertical stems. 

Materials and methods 


From the nature of the experiments, it was necessary to use 
plants with single upright stems. Both woody and herbaceous 
plants were used. The woody plants were both one-year-old 
shoots and seedlings of the common locust {Robin la Pseudo- 
Acacia), both young and one-year-old shoots of sumach (R*** 
glabra), one-year-old shoots of the poplar (Pop id us tremuloides), 
and the castor oil plant (Ricinus communis). The herbaceou 
plants were the sunflower {Helianthus annum), the broad or 
Windsor bean {Vicia Faba), and the common bean (Phascolus 


vulgaris). The experiments upon woody plants were conducted 
in the garden during the spring and summer; those upon her- 
baceous plants were conducted in the greenhouse during the winter 
and in the garden during: the summer. 


After many preliminary trials, the methods given below were 

most suitable. For methods 

tions are given in the account of the different experiments in which 
they were used. In general, a large series of plants of the same size 
and age and growing under the same external conditions was used 
for each experiment. The plants were numbered consecutively. 



part above the compressed region. In case a stem was not cylin- 
drical, the average of the greatest and the least diameter was taken 
as the diameter of the stem at that place. Strong cords were then 
looped about each stem at a suitable distance above the ground 
to serve as an attachment for the weights. The stems were pro- 
tected from mechanical injury by the cords by first wrapping 
pieces of soft cloth, commonly known as table matting, around 
each stem. Each stem was then firmly fastened in a vertical 
position by tying it to stakes, not in contact with the plant, so 
that it could not bend or sway, and yet could be restrained in no 
way except by the downward pull of the weights. Weights were 
then suspended upon half of the plants, while the other half were left 
to serve as controls. All plants in a series thus treated were under 
exactly the same conditions, except that the lower part of each 
experimental plant was subjected to a compression caused by the 
downward pull of the weights. If the plants were practically 
equal in size, alternate stems were weighted; but if they happened 
to be somewhat unequal, care was taken to see that the average 
size of the experimental plants was equal to the average size of 
the control plants. For every large experimental plant, an equally 
krge control plant was chosen, and for every small experimental 
Ptant, an equallv small control olant was chosen. 

More weights were added from 





condition of the several experiments. In some experiments the 
weights were light, scarcely exceeding the weight of the plants; in 
other experiments they were equal to many times the weight of 
the plants themselves. Great care was taken to see that the stems 
were not bent by the weights. 


At the conclusion of each experiment, all the plants were meas- 
ured in the same manner as at the beginning, and the stems taken 




1'IG. I 



compressed part of each experimental plant, as well as the corre- 
sponding part of each control plant, was tested for rigidity- as 
shown by its resistance to bending, and for resistance to crushing- 

The resistance to bending was determined by a simple piece o 

.... 6 - ^ The 

apparatus, which is represented by the 


(fig- i). 



cord was looped over the stem at b, always at a fixed distance 
from a; the weight w was then slowly released and the amount 




of bending read in degrees upon the arc of the protractor mn. 
The testing weight for each series of plants was selected so that the 
plants would not be broken or too strongly bent. The pulley p 
could be adjusted as indicated by dotted lines, in order that the 
weight should at all times act in the direction perpendicular to the 
stem. This method was found to show very slight differences in 
the rigidity of stems. 

The resistance to crushing was determined by the apparatus 
shown in fig. 2. A piece of the 


mm. in length was 

fixed block /*, and the movable 


block m brought against the opposite end. The 

attached to the strong spring balance s, which in turn was attached 

1 * 





- A _ x » ' ' a ' * ' ' 

r* i*o f0 





Fig. 2 

to the threaded rod r . When the tail nut b was turned upon the 
rod r, the movable block m was drawn toward the fixed block f, 
thus subjecting the stem st to pressure. The amount of pressure 
could be read at once upon the scale of the balance. The resistance 
to crushing could be determined very sharply, for when the critical 
point was reached, the stem always gave way suddenly. 

After testing a piece for resistance to crushing, the part of the 
stem next to that which was crushed was put into alcohol to be pre- 
served for a study of the internal structure and microscopic meas- 
urements of the mechanical tissues. 


The anatomical studies and measurements were made from cross- 
sections, which were suitably prepared and stained so as to differ- 
entiate more, clearly the various tissues, principally the xylem 
and hard bast, Chemical reactions (aniline sulfate with sulfuric 
acid, and phloroglucin with hydrochloric acid) as well as stains 
(acid fuchsin, aniline safranin, and methyl blue) were used. 

262 BOTANICAL GAZETTE ' [october 

The method which gave the best results and which was generally 
used was as follows: Cross-sections of uniform thickness were 
placed in watch glasses with a little 50 per cent alcohol and two 
or three drops of alcoholic safranin. In the course of three or four 
days, all the lignified cell walls were stained a bright red, while 
other tissues remained nearly colorless. The sections were then 




Since xylem forms by far the greatest part of the strengthening 
or mechanical, tissue, more measurements were made of this than 
of the other tissues. For this purpose, exact camera drawings 


determined by means of a polar planimeter. Thickness of wall 
was determined by taking the average of several measurements, 
which were made by means of an eyepiece micrometer. Relative 
size of cells was determined by counting the number in a certain 
field in the microscope or by micrometer measurements. 

In order to avoid the danger of a personal error in making the 
mechanical tests and microscopic measurements, the stems were 
numbered indiscriminately by a second person, so that the writer 

xperimental and 
id measurements 

results copied. 



In April, before the growth had begun, 14 one-year-old plants 
re selected from a large number of sprouts. These plants were 





size. More weight was placed upon the experimental plants 

In October 

from time to time, as long as they continued to grow. 

when the experiment was concluded, some of the larger stems W< 

supporting a weight of 20 kilos. 

The results for this series of plants were not satisfactory in a 
respects. Several of the stems were broken or otherwise injure 



during the summer. The growth was very unequal in different 
stems. Some of them increased in size three or four times as much 
as others, which grew under apparently the same conditions. 
This is not an uncommon occurrence, however, among shoots 
which spring from old roots. 

Careful comparisons did not show any greater increase in growth 
or thickness of stem in the experimental plants than in control 
plants. On the other hand, some of the experimental stems had 
smaller diameters in the compressed portion than control stems. 
This diameter in three stems was less than the diameter of the 
same stem above the compressed part. Although this difference 
is normally not infrequently noticed in this species, it was rather 
more striking than usual in these plants. Cross-sections were 
prepared from experimental stems, control stems, and stems which 
had not even been tied to stakes. No differences, however, could 
be seen between any of them. 


In order to obtain woody stems of more nearly uniform size, 
seedlings of locust w r ere grown in the greenhouse and transplanted 
into the garden as soon as the weather became favorable in the 
spring. Of about 100 which were transplanted, only 30 were con- 
sidered suitable for experimentation. These were prepared in the 
usual way and kept under experimental conditions until growth 
had ceased, about October 15. In the greenhouse, some of the 
seedlings grew to a height of 1 5-20 
very weak. When they were transplanted, they were bent nearly 
if not quite to the ground by every little breeze, which could scarcely 
bend locust sprouts which had come up in the garden. In two or 
three weeks' time, however, these slender seedlings, although they 
had grown but little in height, had become strong enough to with- 
stand ordinary winds. Some of these slender seedlings were tied 


wind. Two 

or three weeks later, when the stakes were taken away, the plants 
were still unable to support themselves against a light wind. 

This series was kept under the best conditions for growth. The 
plants were watered frequently during the dry season and protected 


from all injury. The final measurements show that the plants 
grew well. The experimental plants increased 152 per cent in 
height, 131 per cent in the diameter of the weighted portion, and 
122.5 P er cent in the diameter a short distance above the weighted 
part; the control plants, in the same positions and dimensions, 

142 per cent, 135 per cent, and 133.7 per cent respectively. The 
relative amounts of xylem for the experimental and the control 
plants were 95 . 17 and 100 respectively. Careful measurements of 
cell walls showed no difference between the experimental and the 
control plants. 


From a very large number of thrifty sumach sprouts 24 one- 
year-old plants were selected for experimentation. In April these 
plants were tied to stakes, and weights were placed upon the experi- 
mental plants in the usual manner. Conditions were fairly good 
during the growing season. The plants were kept under expen 



in size and thickness of stem took place in the spring and early 
summer. The final measurements and tests showed that the 
experimental and the control plants were practically equal in size 



about 18 per cent less for the experimental than for the control 
plants. Careful examinations and measurements showed no differ- 
ence in the size of cells or in the thickness of cell walls. 


O \J 1SXXVV. II \ 1UU1>U XT^ X w>y 

Sixteen young plants were placed under experimental condition 
as soon as they had attained a height of about 35 cm. These plants 
did not increase greatly in size and thickness of stem. The tests 

aim measurements snowed that the strength ana amount ^* -,- 
were very nearly equal for the experimental and the control plants. 
The percentage increase was somewhat greater for the experimenta 
plants than for the controls. This difference, however, loses I s 
significance when we observe that the actual increase in height wa 

1.- * . r.„ .1 . . , _„ r fh* rnntrol 


plants, and the increase in diameter only 2.07 mm. and 1.7S 



respectively. Careful microscopic examinations and measurements 
failed to show any differences in cells or cell walls. 


Sixteen plants of the poplar were selected and placed under 
experimental conditions in April. The plants were measured and 
carefully tied to stakes. Weights were suspended upon eight of the 
plants, and as the plants increased in size more weights were added. 
Although growth practically ceased in early summer, the plants 
were kept under experimental conditions until October. 

The final measurements and tests showed no greater increase 
in the experimental plants than in the control plants; the increase, 
in fact, was somewhat less for the former than for the latter. For 
the experimental plants, the increase in height, and diameter of 
the stem was less than 16 per cent, and for the controls 20 per cent; 
the relative increase in xylem 77 and 100 respectively; the resist- 
ance to bending 81 and 100 respectively. It is difficult to account 
for these differences. Although the conditions for growth were not 
good, owing to partial defoliation by insects, they were the same 
for both experimental and control plants. 


Twenty-four plants of the castor oil plant were selected from a 
large number which were grown in the greenhouse. Before tying 
the plants to the stakes, the stems were carefully protected with 
pieces of table matting. Internodes of bamboo, about 10 cm. in 
length, were split in two and the halves bound upon opposite sides 
of each stem, the lower ends extending to within 5 cm. of the ground. 
Weights were suspended from the upper ends of these pieces of 
bamboo. In this way room was made for the weights to swing 
free from the ground, and the longitudinal pressure was confined 
to the lower 5 cm. of the stems. The plants were kept under 
experimental conditions for four weeks. The stems 
in diameter and 1 =c cm. hieh at the beginning of the 


3 cm. high at the beginning of the experiment; 
they increased about 50 per cent in diameter and 70 per cent in 
height during the experimental period. At the conclusion of the 
experiment, the experimental stems were under a pressure of 



The results of the final measurements and tests are shown by 
percentages. Assuming in reference to each measurement and test 
that ioo represents the value for the control plants, we have for 
the experimental plants the following numbers : height at the begin- 
ning of the experiment 97.67, at the conclusion 92.33; diameter 
at the beginning 100.2, at the conclusion 104.5; resistance to 
bending 120.6; resistance to crushing 104. 2. The slightly greater 
bending and crushing resistances w r ere probably due to the some- 
what greater diameter of the experimental plants. 

Cross-sections of the stems in this series were prepared and 
studied with special reference to the hard bast fibers and the col- 
lenchyma. The number of hard bast fibers in each stem was 
ascertained by counting. The average number for the experi- 
mental plants was practically equal to the average for the control 
plants. In each stem the thickness of three collenchyma walls 
was measured by means of an eyepiece micrometer, and from these 
measurements the average width of the collenchyma walls deter- 
mined; there was found to be a difference of only 0.07 p between 
the two averages; moreover, this excess was on the side of the 
control plants. 


Eighteen castor oil plants were selected from a large number 
grown in the garden. These plants were tied to stakes and weighted 
in the usual manner. They were about 1 7 cm. in height and 9 . 5 mm ' 

diameter at the beginning of the experiment; they increased 



experimental period of ten days. At the conclusion of the experi- 
ment, the experimental plants were supporting 5-7 kilos eac ' 
The usual measurement and test showed no marked differences. 
Letting 100 represent the value for each control plant average, the 
following numbers represent the relative values for experimenta 
plant average: height at the beginning of the experiment ioi.3> 
at the conclusion 97.2; diameter at the beginning 10 1.6, at t e 
conclusion 101.4; resistance to bending 101.8, to crushing i<>7-5> 

amount of xylem 109 . 3 
Another experiment 



plants, which were divided into three equal sets: experiment plants, 
control plants, and counterbalanced plants. Each stem of counter- 
balanced plants was not only tied to prevent bending, but relieved 
from supporting the weight of the plant itself by tying a cord about 
the stem, passing it up over a pulley, and attaching a weight to the 
free end; this weight was kept practically equal to the weight 
of the plant. These plants were about 2c 
in diameter at the beginning of the experiment; during the experi- 
mental period they increased about 40 per cent in height and 22 
per cent in diameter. The growth was less in this experiment than 
in the former experiment because of more unfavorable weather 
conditions. The final measurements and tests showed no more 
marked differences than in other experiments. 

cm. in height and o mm 


From plants of the sunflower which were grown in pots in 
the greenhouse, 30 of those with straight stems of nearly uniform 
diameter were selected for experimentation. The stems for this 
experiment also were bound upon opposite sides with pieces of 
bamboo to localize the longitudinal compression in the lower 
portions. More weights were suspended upon the pieces of bamboo 
from time to time, until at the end of the experimental period of 


230 grams each. 


about 5.25 mm. at the conclusion of the experiment. Assuming 




control plants, the following numbers represent the values for cor- 
responding measurements and tests in the experimental plants: 
height at the beginning of the experiment 99 . 8, at the conclusion 
97-45 diameter at the beginning 95.9, at the conclusion 98.4; 
resistance to bending 66.7, to crushing 93.5. These numbers 
show very clearly that there was no greater increase in the experi- 
mental than in the control plants. The test for resistance to 

^vv*M-i^ wJA, 

in the control plants. 
Although the mec 






tion made to determine the relative amounts of collenchyma, 
hard bast, and xylem. In none of these tissues could any difference 
be detected between the experimental and the control plants. 


er ulants were selected from ; 

plants and placed under experimental conditions in June. Longi- 
tudinal compression was confined to the lower part of the stems by 
using pieces of bamboo, as in former experiments. Conditions 
for growth were very favorable. The plants were about 15 cm. in 

ght and o mm 

during the experimental period of nine days, they increased 100 per 
cent in height and 70 per cent in diameter. The weights, which 
were necessarily light at first, were increased daily until the average 



experiment the usual mea 

and tests, with one exception, were made. Instead of determining 
the resistance to crushing in this series, the tensile breaking- 

determined bv means of Bordner 

If we assume that 100 


and test for the control plants, the following numbers represent 





conclusion 99.4; resistance to bending 80; tensile breaking- 
strength 64.2; amount of xylem 80.4. These numbers show 

that, although the amount of growth in both the experimen 
and control nlants was nrartimllv mnal thp train in strei 


o — 

in mechanical tissue was less in the former than in the latter. The 


the production 


:ond series of 24 somewhat older sunflower plants was place 
xperimental conditions in the usual manner by tying t e 


plants to stakes and suspending weights upon half of the stems. 
The plants were 22 cm. in height and 8 mm. in diameter at the 
beginning of the experiment; they increased about 140 per cent in 
height and 50 per cent in diameter during the experimental period. 
The final measurements and tests showed very little difference 
between the experimental and the control plants. 

A third series of 81 selected sunflower plants was divided into 
three sets and placed under experimental conditions. The first 
set consisted of 29 experimental plants, the second set of 29 control 
plants, and the third set of 23 control plants which were relieved 
from supporting their own weight by cords looped around the stems, 
passed over pulleys above the plants, and attached to weights, 
which were kept sufficiently heavy to counterbalance the weight 
of the stems. The plants in this series were about 38 cm. in height 
and 10 mm. in diameter, and were kept under experimental con- 
ditions ten days. During that time they increased 80 per cent in 
height and 35 per cent in diameter. Weights were added daily 
until each experimental plant supported 6-7 kilos. 

The usual measurements and tests showed a very great simi- 
larity in every respect in all three sets. In no case was there a dif- 
ference of more than 6 . 1 per cent between the averages for any two 
sets. The relative resistance to bending was as follows: experi- 
mental plants 106. 1, control plants 100, counterbalanced controls 
96.2. The relative amount of xylem was: experimental plants 
103; control plants 100; counterbalanced controls 98. 


Several experiments were conducted in the greenhouse with 
the Windsor bean. Experimental plants tied to stakes and with 
the longitudinal compression localized in the lower portion, control 
plants tied to stakes, counterbalanced controls, and plants which 
grew normally, without additional weight or support, were com- 
pared with reference to strength of stem and amount of mechanical 
tissue. In no case did additional weight or decreased weight seem 
to have any influence upon the plants. Stems of plants which 
had been tied to stakes were found to be slightly weaker than 
stems which had not been tied. 



Twenty-four plants of the common bean were selected and 
placed under experimental conditions. The longitudinal com- 
pression was localized in the lower parts of the stems by means 
of the pieces of bamboo. Measurements and comparisons showed 
no greater gain in mechanical tissue in the experimental plants 

than in the control plants. 



special reference to compression may be considered under five 
heads: (1) heredity and variation; (2) correlation; (3) mechanical 
effects of compression; (4) stimulatory effects of compression; 
(5) compression in connection with other stimuli. 


The production of a certain amount of mechanical tissue in 
plants is a specific characteristic just as much as is the production 
of leaves of certain form. Just as leaves of a given species may 
vary in number and size under the laws of fluctuating variability, 
so may the amount of mechanical tissue vary. Entirely independ- 
ent of any known cause, sudden variations may occur in the amount 
of mechanical tissue formed. The best illustration of this sudden 
variability is furnished by Oenothera lata (De Vries 9), a mutant 
which does not produce enough mechanical tissue to support its 
own weight. Another common illustration is furnished by certain 
laciniate-leaved varieties of the sumach, which have stems too 
weak to bear their own weieht. The difference in the amount 

s or 

of xylem in closely related forms is shown in many grafts. When 
a scion of one species of elm is grafted upon another specie 
variety, it soon becomes larger than the branch upon which it was 
grafted, although there is more mechanical strain upon the branch 
than upon the scion. The same phenomenon is frequently seen in 
grafts upon fruit trees. 

All of my experiments, especially those in which the weight 
the plants was counterbalanced, showed that it is characteristic 
of each of the species to produce a certain amount of mechanica 


tissue. It may be that at some time in the history of a plant it 
was susceptible to mechanical stresses, and that as a result of these 
stresses the plant gradually came to develop normally a certain 
amount of mechanical tissue without regard to the stresses to 
which it might be subjected. Whether this is the case or not is a 
mere matter of speculation, so far as this work is concerned, since 
of course experiments can throw no light upon that phase of the. 
problem. Yet some plants may still be able to respond in a slight 
degree to such stimuli. It was with this possibility in mind that 
such comparatively large numbers of plants were used and careful 
measurements made in order that individual variations might be 
obliterated in the average of a large number. 

It is possible and even altogether probable that the importance 
of such differences and variations may not have been given the 
proper amount of attention, or that they may have been overlooked 
entirely. In this work, however, the possibility of an error from 
both specific and individual variation has been eliminated by 
getting the averages of a large number of the same species for 
each experiment. Further discussion of this phase of the question, 
therefore, may be omitted. Heredity, however, does have a very 
mportant bearing upon the problem in another way. 

The ability to produce certain tissues may be hereditary but 
latent in a plant, and require a certain definite stimulus to make 
it appear. Many plants do not produce leaves unless they feel 
the stimulus of light; others, cacti for example, produce leaves 
only when deprived of light. , The experiments of Von Derschau 
(35) and Worgitzky (37) show that mere contact may cause the 
production of a certain amount of mechanical tissue, and contact 
and strain together are necessary to cause a tendril to produce its 




ments show that, at least in the plants which were used in this 
work, no latent hereditary activity is brought forth by compression. 


Under this head may be considered all those observations upon 

branches. Den 



ber of instances which they investigated that anatomical changes 
resulting in greater strength go hand in hand with the develop- 
ment of the fruit. These changes have been attributed both to 
the strain which is caused by the weight of the developing fruit, 
and to greater growth activity which is due to the carrying of the 
greater amount of building material which the fruit demands. 
Keller (16) affirmed, however, that the fruit-bearing axis is not 
strengthened by strain or fruit-bearing in itself, but that the 
bending of the stem from the orthotropic to the plagiotropic 
position causes the anatomical changes. It is evident that these 
observations cannot throw any light upon the stimulatory effect 
of a tension or a compression acting alone. An experiment by 
Vochting (34), however, does have a more direct bearing upon 
the question. He found that by removing the flower buds he could 
greatly retard the development of the mechanical tissue; the plants 
were sunflowers and fruiting stems of the Savoy cabbage. He also 
found that by stress brought about by weighting he could not cause 
the mechanical elements to reappear (the discussion of this stress 
is taken up in another place), but when he grafted a scion upon a 
decapitated stem, the cambium at once began to form the mechani- 
cal elements aga'in, although the weight of the scion was equal to 
only a fraction of the weights which he had placed upon similar 
plants without result. He sums up the results of his experiments 
in the following words : 

Aus dem mitgetheilten folgt, dass der ontogenetische Gang der Gewebe- 
differenzierung von inneren, correlativen Verhaltnissen beherrscht, dass die 
Bildung der einzelnen Gewebeformen nicht einfach durch das Bedurfmss 
bestimmt wird. Die Auslosungs-Theorie genugt hier nicht. 

This experiment shows very clearly that correlation may have much 


111 iv vw witii incciuuuuai tissues uimi meic sucsa w* ^ 

Correlation is also seen in many trees in which one-sided growt 
in the wood is correlated with a greater development of the root 
system on that side, or with the greater number of branches an 
the correspondingly greater leaf surface of that side. Knv (ifl 
asserted that one-sided leafing of a tree must b? associated wit 
one-sided increase in the growth of the trunk. Hartig (io) r< 



pine trunks, although there were exceptions to the general rule. 
Wiesner (36) has attempted to explain eccentric growth in 

branches by attributing it to unequal size and distribution of the 


In accordance with the observations given above, it was found 



plant. Locust shoots, which at one time w^ere apparently equal 
and growing under equal conditions, were found to develop at very 
unequal rates, apparently because of differences in the amount of 

nutriment stored in the roots from which the separate shoots had 

The differences in herbaceous stems under different conditions 
are perhaps best illustrated by sunflowers which were grown in the 
greenhouse and plants of the same kind and age which were grown 
in the garden. The tall, slender greenhouse plants at the end of 
four weeks had such poorly developed xylem cylinders that many of 
the stems were not able to support the weight of the tops, while 
the garden plants in the same time had become strong and robust, 
with well-developed xylem cylinders. This is, of course, an extreme 
case. Many plants, however, which were grown under apparently 
the same conditions had to be rejected because of abnormalities 
or differences in growth. By selecting plants which were as nearly 
uniform as could be obtained, and by keeping all conditions of 
growth as nearly equal as possible, it was thought that accurate 
results could be obtained, especially since the possibility of a per- 
sonal error was also excluded. 


The textbooks assert that pressure, or compression, retards 
growth and may completely stop it if the pressure is great enough 
(Peirce 24, Jost 15, Pfeffer 26). It seems no more necessary 
to make this assertion than to assert that heat, for example, may 
retard growth or even kill the plant if- the temperature be high 
enough . It is of importance, however, to know whether retardation 
is proportional to the pressure, and whether changes take place in 
the cell or in its walls. 


The effect upon the plastic growing cell must be the same 
whether it is compressed or whether it meets resistance to further 
growth. There can also be no difference in the hydrostatic or the 
osmotic pressure within the cell, whether the pressure be in one 
direction or another. In stems a longitudinal pressure causes the 
cells to exert a greater lateral pressure upon each other, and vice 
versa a lateral pressure must cause them to exert a longitudinal 
pressure upon each other. 

Earlier writers assumed that growth must be inversely propor- 
tional to the external pressure exerted upon the growing cells. 
According to this view, the greatest growth must be in the region 


of the least pressure. It was upon this assumption that Detlef- 

sen (7), Sachs (30), Nordlinger (23), and Kny (17) sought to 

explain the eccentric growth of stems and branches. They had 
then merely to account for a loosening of the bark in some way 
to cause increased growth. De Vries (8) would account for the 
annual rings by assuming that the bark pressure is greater at some 
seasons of the year than at other seasons. The experiments, in 
which he retarded growth and caused the production of smaller 
cells by putting ligaments around the trunk, showed that great 
pressure may retard growth, but his relieving pressure by cutting 
the bark with a knife produced traumatic effects and not true 
growth. Kr abbe's (18) experiments proved that these theories 
for unequal growth were erroneous, for he showed that the bark 
pressure is never great enough to retard growth. This was an 
indication that growth cannot be directly or indirectly propor- 
tional to the pressure. This point was still further emphasized 
and cleared up by Pfeffer (25). Pfeffer says also (Plant 
Physiology 2:125): 

When a root pushes an object in front of it, its rapidity of growth is not 
perceptibly affected unless the resistance offered is very great. 


the manner in which the cell meets external compression, thi 




The manner 




Under such circumstances [when a plant part is compressed] the tension 
of the cell walls is gradually replaced by the external pressure, against which 
the whole osmotic pressure finally acts. 

It is thus seen that the plant does not meet a compressing force 
by growth and cell division, but by osmotic pressure. In reference 
to growth activity in cells under compression, Noll's (22) experi- 
ment seems very significant. He showed that in bent roots second- 
ary roots do not arise from the concave side where the cells must 
be compressed. 

• Hartig (ii), Cieslar (5), and Sonntag (31) attributed the 
increased thickness of cell walls in heartwood to pressure. Now 
it does not seem reasonable to think that pressure could cause the 
walls to become thicker, unless it acts as a stimulus merely to cause 
greater protoplasmic activity. Since the plant opposes external 
compression, or resistance to growth, by its osmotic pressure, it 
would be detrimental to the plant to have the walls lignified before 
their limits of expansion had been reached. That secondary thick- 
ening does not take place until the extension of the cell wall has 
ceased seems to be the general rule. Pfeffer (op. ciL, p. 27) says: 

After the stretching growth of the cell has ceased, the cell wall commonly 
undergoes secondary thickening; 

and again (op. cit., p. 32) : 

An intimate correlation exists between the different forms of growth, and 
the thickening, cuticularizing, or lignification of the cell wall usually take 
place when the external growth has ceased. 

In regard to the part which the increase in thickness plays in 
overcoming resistance, we may again quote Pfeffer (op. cit., p. 


Even when external growth has ceased, the thickening of the cell walls 
may still further increase the pressure which the growing tissue is capable of 

exerting. The observed pressures, ho 
produced by the osmotic pressure alone. 


stems that thickening of cell walls was always retarded in the 
parts of the stem which were under compression. 

The best experimental evidence of the effect of pressure upon 
cell walls was given bv Newcombe (20) . Plant stems were inclosed 


within plaster casts; as they increased in diameter, the pressure 
became greater. It was found that under these conditions the 
cells remained alive and active, yet the walls remained thin. Even 
the walls which had already begun to thicken ceased to become 
thicker as pressure increased. 

My experiments in which very young sunflower stems were 
subjected to heavy weights showed fairly well the retarding effect 
of a longitudinal compression, for the amount and the strength 
of the mechanical tissue was found to be less in the experimental 
than in the control plants. 


Very little is to be found in botanical literature upon longi- 
tudinal pressure, or compression, as a stimulus, and still less as to 
its effect or lack of effect upon mechanical tissues. In nearly 
every instance in which the stimulatory effect of pressure is men- 
tioned, it is loosely applied or very evidently connected with other 
causes of stimulation. Pfeffer states (op. cit., p. i24) : 

Various stimulatory influences are exercised by tension, pressure, an 
mechanical vibrations or disturbances. 

What he says concerning the influence of compression has already 
been discussed under mechanical effects. 

In nearly all researches, compression has been associated with 
other stimulatory influences. Those observations in which the 
author attributed the observed effects to pressure, or compression, 
are discussed under this head. 

Hartig (10) found that trees which were subjected to the action 
of a west wind produced thicker annual rings upon the east side 
than upon the west side. His explanation is best given in his 
own words: 

Der Druck des Windes ubt einen Reiz auf das Plasma der Cambiumschicht 
aus, welche in zweckentsprechender Weise durch gesteigertes \\ achsthum un 

durch Dickwandigkeit der Organe auf diesen Reiz reagirt. 

These observations have not been confirmed by experiment 
and Ursprung (32) has shown by his observations and experi 
ments that longitudinal compression cannot account for these 



Cieslar (5) bent four young spruces at right angles so that the 
upper parts lay in a horizontal plane, and kept them bound in 
that position for two years. He found the greatest increase in the 
thickness of annual rings to be in the concave or compressed side. 
From this experiment he concluded that pressure causes the greater 
increase in growth. The objections to this conclusion, however, 
are obvious. Cieslar apparently did not consider the tension 
upon the convex side, nor the one-sided effect of gravity, which 
must produce a geotropic influence as soon as the tree is bent 
from the perpendicular position. The same objections may also 
be made to Sonntag's (31) conclusions. 

Vochting's (34) experiments, however, require more con- 



question whether his plants were placed under longitudinal com- 
pression, tension, or a combination of the two. After finding that 
sunflower and cabbage plants, which had been decapitated to 

prevent fruiting, did not produce the normal amount of mechanical 
tissue, he says : 

Zunachst und vor allem fasste man die mechanischen Zellen in's Auge. 
Nach Hegler's bekannten Angaben kann man durch kiinstliche Belastung 
nicht nur eine raschere Entwickelung dieser Elemente herbeifuhren, sondern 
ihre Xeubildung sogar an Orten bewirken, an denen sie sonst nicht entstehen. 
1st diese Angabe, die durch unsere eigenen Erfahrungen nach gewissern Richtung 
bestatigt wurde, rich tig, dann darf man erwarten, dass die veranderten Objecte 
unseres Versuches, wenn kiinstlich belastet, die mechanischen Elemente 
wieder hervorbringen. Von dieser Voraussetzung ausgehend wurden am 
Bluhen verhinderte Pflanzen des Wirsing und der Sonnenblume durch Gewichte 



Experimente verwandt. Die Belastung war verschieden gross; die grosste 

Kilo, beim Wirsing mehr 

unter gewohnlichen 

nissen von der Pflanze erzeugt werden, um ein Vielfaches iibertreffen. Die 
Belastung dauerte Monate lang. 


experiments in which stems 

Weise" to mean 

stems. This is the interpre- 

(i), HlBB 



that a plant responds to the demands ("das Bedurfniss") made 
upon it, we may interpret the phrase to mean that he fastened 
the weights in such a way as to simulate the tension caused by 
the fruiting head. Bucher (4) interpreted this to mean a longi- 
tudinal pressure or compression, as indeed it must if the fruiting 
heads remained erect. Since, however, the fruiting head of a 
sunflower droops, it must exert tension upon the part of the stem 
next to it, and both tension and compression in the bent part, 
tension upon the upper or convex part and compression upon the 
under or concave part. It seems from a later statement of Voch- 
ting's that the stems were kept erect. In that case, the "Belas- 
tung " must have exerted a compression only. He says in summing 
up his results : 

Wir fanden, dass selbst sehr starke Belastung der aufrechten am Bliihen 
verhinderten Objecte nicht genugt, um das normale Holzwachsthum wieder 


and less cambial activity in the two castor oil plants which he 
subjected to longitudinal compression than in normal plants. 
The stems were also larger in diameter, owing to the abnormally 
large thin-walled cells. 

Hibbard (14) used the same general method for subjecting 
stems to compression that I have employed in this investigation. 
His experiments were of a preliminary nature, comparatively 
few plants were used, and there is no record of his making mechan- 
ical tests or exact measurements of the mechanical tissue. When 
so few plants are used and no averages made, errors from indi- 
vidual variation may easily occur. It is not certain that the 



the same conclusions were arrived at by two persons working 

In reference to Hibbakd's experiments as well as to my own, a 
question may arise as to whether a stimulus is really exerted by 
such means as were used. This question may be answered in 
several ways. It is well known that living protoplasm is very 
sensitive. This sensitiveness is shown by the responses of Mimosa 


leaves and by tendrils. The researches of Bose (3) have shown 
that the protoplasm of other plants and plant parts is equally- 
sensitive, although the plant cannot respond by a perceptible 
movement. In case growth is retarded, as it was in some of the 
experiments, the protoplasm must necessarily feel the effect. It 
has been demonstrated that there are interacting stresses in plants, 
of which a good example is afforded by twisted tree trunks. Force 
is required to cause such a twisting. If the weight of the top is the 
force, twisting must be accompanied by a shortening of the central 
cylinder or by an elongation of the outer layers. In either case, a 
longitudinal compression must be exerted upon the outer cells. 
It is not certain that the inner cylinder may become shorter, but 
it has been shown that wood cells may increase in length from the 
center to the circumference of a tree trunk. If this elongation of 
cells tends to lengthen the outer layers of wood in straight trunks, 
increased weight upon the trunk must alter the internal equilibrium 
of stresses, for it not only acts directly upon the outer layer, but it 
tends to increase the retarding action of the inner cylinder. More- 
over, it is not safe to say that the woody cylinder is so rigid that it 
cannot change, for we know that wood is more or less elastic, and 
that the limit of elasticity may be exceeded by either great pressure 
or great tension. If a bar of wood is forcibly bent and kept in 
that position for some time, a readjustment takes place so that 
the bar remains bent without the external force. The sensitive 
protoplasm in a cambium layer, which surrounds closely a cylinder 
of wood, must feel very slight changes that might occur in the 

If the weight of the plant can furnish a stimulus for production 




srL ^ 

the same reason an increased weight, equal to two, four, ten, or 


twenty times the weight of the plant, ought to produce a response 
by an increase in the strengthening tissue. Assuming that plants do 



experiments were placed under the most favorable conditions to 
respond to a given stress, for they were relieved of all stress except 


the downward pull of the weights. In no case, however, were they 
found to become stronger mechanically. It must be concluded that 
a longitudinal compression, acting by itself and acting continuously 
upon a plant stem, cannot stimulate it to greater activity, nor to 
an increased strengthening of the tissues that are already present. 


For convenience, we may separate the stimulatory influences into 
three groups: the so-called internal stimuli, environmental stimuli, 
and strains or stresses. These internal stimuli may be due indirectly 
to environmental conditions. Since we know little or nothing 
about them, it is convenient to refer to this group all those phenom- 

ena which we cannot attribute to definite external conditions or 
stimuli. Environmental stimuli include gravity, light, heat, 
moisture, etc. Strains or stresses are tension and compression. 

In nature the various influences are intimately associated with 
each other, and often so modify one another that observers and 
sometimes experimenters have overlooked certain influences and 
thus given undue weight to others. As an example we may speak 
of Hartig's observations and Cieslar's experiments in which 
pressure only was considered, where evidently gravity, tension, and 
perhaps other influences may have played an important part. 

Others, who were not able to account for observed phenomena 
by certain external stimuli, explained them by saying that they 
are due to internal stimuli. As an illustration, a quotation from 
Massart (19) will serve very well. In speaking of root swellings 
and eccentric growth in Ficus roots he says: 

L'epaississement asymetrique est re'gi par la gravitation, par la lumiere e 
par des excitants internes, encore indetermines. 

Experimenters who examine more closely into the reactions 
of plants are forced more and more to the conclusion that t 
plant acts as a unit in a self-regulatory manner to bring about t e 
greatest good to itself. Newcombe (21) was the first to point 
out the necessity of considering the self -regulatory power of plan 
with special reference to their response to stress. UrspR un 
(32, 33) came to practically the same conclusion after a series 
observations upon eccentric growth in trunks and branches. 


Bucher (4) also arrived at this conclusion as a result of trying 
to bring his results into harmony with Ball's (i) conclusions. 

The application of the self-regulatory theory may seem to serve 
merely as a term to cover our ignorance. In reality it throws much 
light upon the whole question of stimulatory reactions, since it 
shows the great number of possibilities. In order to determine the 
influence of one factor, all the others must be taken into consider- 
ation, and their influence either eliminated or equalized by the use 
of both experimental and control plants under exact experimental 

Considerable investigation has been carried on with respect to 
the various environmental stimuli. Further discussion of them, 
however, is not necessary in this place. In regard to the stimula- 
tory effect of stresses and strains, comparatively few exact data 
have been obtained. 



as well as from my own experiments. Bucher found that when a 
stem is bent and then placed upon a klinostat the cell walls on the 
convex or tension side become thicker, and on the concave or 
compression side the walls remain thin, while the cells themselves 
become larger. In stems which were held in a horizontal position 
so that they could not bend upward under the stimulation of 
gravity, he observed the same results; the upper or stretched side 
had normal or slightly smaller cells with thickened walls, while 
the lower side had large thin-walled cells. Now Bordner's results 
show that, under proper experimental conditions, tension causes 
a strengthening of stems by producing thicker walls; and it is 
shown that compression, when not too great, causes large thin- 
walled cells. It 

sion may be given as the two direct influences for the results 
observed by Bucher. 

It is evident that the tension and compression do not have 
exactly opposite effects. We have seen that compression causes 
the cells to become larger and thus give greater surface for the 
osmotic pressure. Tension, as shown by the experiments of 
Noll and Hpt-tttt? t^nHc tr> nrrfApratp <rrowth and cell division. 



The two influences of tension and compression are thus both 
favorable to increase in the size of the organ subjected to them. 


may cause the walls to thicken 


and strength in the part subjected to those stresses. It does not 
seem unreasonable to assume that intermittent or alternating 
compression and tension should have a greater stimulatory effect 
than a constant stress. It is very possible that under constant 
stress the internal equilibrium of stresses may come in time to be 
readjusted so that the external stress would be no longer felt. In 
experimental work this objection may be removed in part by increas- 
ing the stress from time to time. Yet it is easy to see that this 





in mechanical strength or in the amount or kind of mechanical 
tissue under the influence of longitudinal compression. 

In young herbaceous stems the development of mechanical 
strength in the tissues is somewhat retarded by a longitudinal 
compression caused by comparatively heavy weights. 

Neither light weights nor heavy weights have any appreciable 
effect upon the growth and strength of herbaceous stems which 




compression, excepting of course mech 


The investigation for which these results were obtained wa 
carried out under the direction of Professor F. C. Newcombe, 
to whom I wish to acknowledge my sincere thanks for his kindly 
interest and timely suggestions. 

University of Michigan 
Ann Arbor, Mich. 



1. Ball, 0. M., Der Einfluss von Zug auf die Ausbildung von Festigungs- 
gewebe. Jahrb. Wiss. Bot. 39:305. 1904. 

2. Bordner, J. S., The influence of traction on the formation of mechanical 
tissue in stems. Box. Gazette 48:251. 1909. 

3- Bose, J. C, Plant response. London. 1906. 

4- Bucher, H., Anatomische Veranderungen bei gewaltsamer Krummung, 
und geotropischer Induktion. Jahrb. Wiss. Bot. 43:271. 1907. 

5. Cieslar, A., Das Rothholz der Fichte. Centralbl. Ges. Forstwesen, 
p. 149- 1896; quoted by Kny, L., Jahrb. Wiss. Bot. 37 : 62. 1896. 

6. Dennert, E., Anatomische Metamorphose der Bliithenstandaxen. Bot. 
Heft. Forsch. Bot. Gart. Marburg 2:128-217, 1887; abstract in Bot. 
Jahresb. 15:647. 1887. 

7- Detleesen, E., Versuch einer mechanischen Erklarung des excentrischen 
Dickenwachsthum verholtzer Achsen und Wurzeln. Arbeit Bot. Inst 



des couches des ligneuses annuelles. Archives Neerlandaises 11: 

9« ~ -, Die Mutationstheorie. Leipzig. 1900. 

10. Hartig, R., Holzuntersuchungen. Altes u. Neues. Berlin. 1901. 

11 • ; Das Rothholz der Fichte. Forst. Nat. Wiss. Zeit. 1896. 

12. Hegler, R., Ueber den Einfluss des mechanischen Zugs auf das Wachs- 

tum der Pflanzen. Cohn's Beitr. Biol. Pflanz. 6:383. 1893. 

x 3« ~ , Reported by Peeffer, q. v. 

14* Hibbard, R. P., The influence of tension on the formation of mechanical 

tissue in plants. Bot. Gazette 43:361-382. 1907. 
15- Jost, L., Plant physiology, English translation. Oxford. 1907. 


Fruchtstielen. Inaug. Diss. Kiel. 1896. 

17- Kxy, L., Ueber das Dickenwachstum des Holzkorpers. Berlin. 1882. 
18. Krabbe, G., Ueber das Wachstum des Verdickungsringes und der jungen 
Holzzellen in seiner Abhangigkeit von Druckwirkungen. Abhandl. 

Mem. Acad. 


Berliner Akad. 1884. 

Massart, J., Sur Tirritabilite des plantes s 

Belgique 62:50. 1902. 

Newcombe, F. C, The influence of mechanical 

ment and life period of cells. Box. Gazette I9 : i49> i 9 1 ^ 22 9- l8 94- 

20:441. 1895. 


Bot. Gazette 

22. Noll, F., Thiel's Landw. Jahrb. 29:361. 1900; quoted in Bonn Text- 
book. 1908. 


23. Nordlinger, H., Ovale Form des Schaftquerschnittes der Baume. 
Centralbl. Ges. Forstwesen. 1882. 

24. Peirce, G. J., Plant physiology. New York. 1903. 

25. Pfeffer, W., Druck- und Arbeitsleistung durch wachsende Pilanzen 
Abhandl. Konig. Sach. Gesells. Wiss. 20:233. 1893. 

26. , Physiology of plants. English translation, vol. 2. Oxford. 1903. 

27. , Untersuchungen R. Hegler's iiber den Einfluss von Zugkraften 

auf die Festigkeit und die Ausbildung mechan. Gewebe in Pflanzen. Bcr. 
Kgl. Sach. Gesells. Wiss. Math.-Phys. CI. 43:638. 1891. 

28. Pieters, A. J., The influence of fruit-bearing on the development of 
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30. Sachs, J., Lehrbuch der Botanik. 1873. 

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turns. Biol. Centralbl. 26:257-272. 1906. 

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Flora 69:2-91- 


Lincoln Ware Riddle 

(with nine figures) 

The American species of Stereocaulon fall naturally into two 
groups. One is a typically boreal group, with S. paschale the 
central species, and a general distribution throughout British 
North America and the northern United States, as far south as 
North Carolina in the Alleghany Mountains, and in the Rocky 
Mountains extending into Colorado, and in the case of some of the 
species even into the mountains of Mexico and the South American 
Andes. The other group is a tropical one, represented by 5. 
ramulosum and its allies, and characteristic of tropical America, 
from which it extends rather widely into the southern hemisphere. 
No American herbarium contains sufficient material for a satis- 
factory study of these tropical species, and for that reason the 
present paper is confined to a consideration of the boreal species. 

Anyone who has attempted to determine material of Stereo- 
caulon, or has studied the material in our herbaria, must have 
realized the confusion which exists in regard to the distinctive 


characters of the species. This confusion arises in part from the 
great variability of the species, and their tendency, in some cases, 
to intergrade. But another cause of the confusion comes from the 
lack of literature available to the general student and containing 
comparative notes on the species. This lack of comparative notes 
makes Tuckerman's Synopsis of the North American Lichens, which 

s been the guide for most of the study of the American material, 
a difficult treatment to use. The classic work on the genus Stereo- 
caulon is to be found in the papers of Th. M. Fries, his De Stcreo- 
cauhs et Pilophoris commentalio being published in 1857, and his 
Monographic Stereocaulorum et Pilophoronim in 1858. These must 
form the basis for any careful study, but unfortunately they are 
rather inaccessible to American students. 

The present paper is an attempt to clear up some of the existing 

- 8 5] [Botanical Gazette, vol. 50 


confusion and to provide an available account of the North American 
species of the boreal group, which shall include comparative notes, 
and shall cite the specimens to be found in some of our herbaria. 
I have been fortunate in having access, through the courtesy of 
Professor Farlow, to the Tuckerman Herbarium, which contains 
authentic material of the following species: 

S. coralloides Fr., S. denudatum Flke., S. nanodes Tuck., S. paschale var. 
conglomeratum Fr., S. tomentosum Fr., S. Wrightii Tuck., S. cereolinum 
Koerb., S. Despreaultii Del., S. glaucescens Tuck., and S. tenellum Tuck. 

Professor Farlow's own herbarium contains an authentic 
specimen of S. foliiforme Hue; and Professor Fink has kindly- 
communicated a specimen of S. alpinum Laur. from Laurer s 
herbarium, which I have taken to be authentic. 

In addition to the Tuckerman collection (cited as "Tuck")> 
the following herbaria have furnished the basis for study. The 
abbreviations in parentheses are those used in the body of this 
paper in the citation of specimens. 

i. Cryptogamic Herbarium of Harvard University, including 
the personal collection of Professor W. G. Farlow. (H) 

2. Herbarium of Wellesley College. (W) 

3. The Clara E. Cummings lichen collection. Wellesley 
College. (CEC) 

4. The C. J. Sprague collection at the Boston Society of Natural 
History. (BSNH) 

5. Herbarium of the Thoreau Museum of Natural History, 
Middlesex School, Concord, Mass. (Th) 

6. Herbarium of the University of Vermont, including the col- 
lections of C. C. Frost and Dr. C. G. Pringle. (UVM) 

7. Herbarium of Brown University. (B) 

8. Herbarium of Yale University. (Y) 

9. Herbarium of the New York Botanical Gardens. (NY) 
10. He barium of the Geological Survey of Canada. (Can) 

Rand). (Mt.D) 

12. Herbariun 



Herbarium of L. W. Riddle. Welleslev. Mass. (R) 

If my interpretation of the principles of generic nomenclature 


is correct, Acharius should be cited as having definitely estab- 
lished the genus Stereocaulon in Methodus Lichenum (1803), p. 314, 
with Lichen ramulosus Swartz as the type species. It is true that 
the name had been used by three previous authors, but none of 
these can be held to have established the genus, according to present 
ideas. Schreber first published the name with a brief diagnosis 
in Linnaeus' Genera Plantarum, ed. 8 (1791), p. 768, but no species 
is cited under the genus. Schrader, in Spicileginm Florae Ger- 
manicae (1794), p. 113, used the name with a single species, 5. 
corallinuni; a species based, however, on an imperfect lichen which 
later proved to be a Pertusaria. Finally, Hoffman, in Deutsch- 
lands Flora, II (1795), p. 128, gives a genus Stereocaulon, with nine 
species; of these nine only three are now recognized as belonging 


is: Hoffs 

genus, therefore. " embraced elements altogether incoherent " and 
hence cannot be considered valid. Acharius thus remains the 
first author to place the genus on a solid foundation. The generic 
characters may be stated as follows: 

Thallus of two parts, a primary horizontal thallus, which in most cases 
disappears, and erect, solid, rvlindrical. ecorticate oodetia: thallus of 


gularly flattened squamules, which are typically 

sometimes creamy or white, and which also cover the podetia more or less 
thickly; apothecia lecideine; spores fusiform to acicular, hyaline, plurilocular. 
Closely related to Cladonia, from which it differs in the solid podetia and the 
plurilocular spores, the latter character also serving to .distinguish this genus 
from Pilophorus. 

The term "squamules" (used by Acharius, E. Fries, Schaerer, 

and others) is here adopted, in preference to the term "phyl- 
locladia" (used by Th. Fries and by Tuckerman) , for the thalline 
outgrowths which constitute the thallus of species of the § Pro- 
stereocaulon, and which occur so characteristically on the podetia 
of all species. The term is in current use for such thalline structures 


in the genus Cladonia, 

with which the squamules of Stereocaulon are strictly homologous, 
although somewhat modified. 

In the descriptions of the species I have made no attempt to 



but rather to give a diagnosis of the salient and characteristic 

wx y v, t*. wiu G 

features by which the species is to be recognized. 


This key is based on typical specimens. For the determination of inter- 
grading forms, reference should be made to the comparative notes, which 
follow each species. 

A. Plants chalky-powdery 9- s - dHam 

AA. Plants not chalky-powdery. 

B. Primary thallus present. 

C. Spores 4/x wide, ends blunt; podetia frequently ending in 

soredia i. S. pileatim 

CC. Spores 2.7 m wide, ends pointed; podetia never ending in 

soredia 2. S. condensation 

BB. Primary thallus absent. 

C. True squamules absent, podetia ending in flattened, foliose 
tips 8. S.Wrightii 

CC. True squamules present. 

D. Squamules of the palmate -digit ate type (see fig. 7)- 

E. Podetia glabrous or faintly tomentose, cephalodia 

with Stigonema. 
F. Podetia loosely branched and spreading 

4. S. paschale 



FF. Podetia in compact cushions 

4 a. 5. paschale v. conglomeratic 
Podetia more or less densely tomentose, cephalodia 
with Nostoc. 

F. Podetia repeatedly branched 
* s . S. tomcntosmn 

FF. Podetia subsimple 

5a. 5. tomcntosum v. simplex 

DD. Squamules of some other type. 

E. Squamules coralline 3- S. coralloidcs 

EE. Squamules umbilicate or coarsely granular. 

F. Podetia glabrous, cephalodia with Stigonema 

7 . S. denudation 

FF. Podetia tomentose. cephalodia with Nostoc 

6. S. alpine 

Section Prostereocaulon 

Primary thallus persistent and closely adnate to the substratum . 
podetia mostly short, under 2 cm., and simple or sparingly branched ; 


squamules poorly developed, mostly granular. — The species of this 
section show resemblance to Pilophorus. 

Koerber's name "Cereolus" for this section must be abandoned, with 
the exclusion of the Lichen cereolus of Acharius from the genus (see dis- 
cussion under S. pileatum). 

1. Stereocaulon pileatum Ach. 

5. pileatum Ach. Lich. Univ. p. 582. 18 10. 

S. condensatum Laurer in Fries Lich. Eu. p. 203, in part. 183 1; and in 
Tuck. Syn. Lich. New Eng. p. 46, in part. 1848. 

S. cereolus Schaerer Enum. Crit. Lich. Eu. p. 178. 1850; not Ach. Meth. 
p. 316. 1803. 

5. cereolinum Koerber Syst. Lich. Germ. (1855) p. 14. 1855; not Ach. 
Syn. p. 285. 1814. 



persistent, closely adnate to the substratum, 
squamules which tend to become coralline; 


with scattered coralline-granular squamules; apothecia terminal, 
or often absent and the podetia then ending in a mass of white 
soredia: spores 16.5-29X3.5-5 Z 4 , average 21.4X3-9^ w * tn 

blunt ends; 
on rocks. 

paschale (q.v.). — Growing 


Europe: Stenh. Lich. Suec. no. 85; Moug. and Nestl. no. 947; Koerb. 

Lich. sel. Germ. no. 271. 

Waghome 1893, as s - nanodes (Hb. Eckfeldt); 

898 (Can); Nova Scotia, Macoun 

Macoun July 


Manan Is., Wilky (BSXH). 



New Hampshire : White Mts., Tuckerman Lich. Exs. no. 113 in part; 
also Tuckerman in Herb.; Bald Mt., W. G. Farlow Sept. 1905 (H); Chocorua, 
W.G. Farlow Sept. 1907 (H); Fitzwilliam, R. H. Howe, Jr., Aug. 6, 1909 (R). 

Vermont: Mt. Mansfield, C. G. Pringle no. 197 (BSNH); Willoughby, 
G. G. Kennedy July 28, 1898 (H). 

Massachusetts: Chelmsford and Salem, /. L. Russell 1850 (Tuck) ; Con- 




This is the only species treated in this paper of which the synonymy is 

2 go 



confused, the uncertainty arising on account of the doubt as to what Acharius 
meant by his Lichen cer coins, described in Lichenographia Suecicae Prodr omits 



S. cereolns being placed as a variety under S. cereolinum (Ach.) Koerb., of 
which S. pileatum is made a synonym, apparently disregarding priority. 
But in his Lichenographia Scandinavica (1871), p. 55, Th. Fries states that 
according to the original specimens in Acharius' Herbarium, Lichen cereolus 
is synonymous with Pilophorus Fibula Th. Fr. This would leave S. pileatum 
the oldest name and the one to be adopted, as has indeed been done by Th. 
Fries {pp. ciL) and most recent authors. 

Stereocaulon pileatum is closely allied to S. condensatum, with which, as 
indicated in the synonymy, it was at one time united by certain authors, yet 
the differences between the two are greater than those between 5. paschale 
and S. tomentosum, which have been recognized as distinct since the work of 
Elias Fries. The points of difference are as follows. 

S. pileatum 

1. Squamules typically coralline. 

2. Podetia simple and reduced. 

S. condensatum 

1. Squamules granular. 

2. Podetia stouter, often branched 

. above. 

3. Apothecia often replaced by sore- 3. Apothecia never replaced by sore- 



4. Spores with blunt ends, and aver- 4. Spores with pointed ends, and 

aging 4 m in width. 
5. Growing on rocks. 

averaging 2 . 7 M in width. 
5. Growing on soil. 

S. pileatum also shows a strong resemblance to forms of Pilophorus. 
When fertile the spore differences will of course serve for their distinction. 

2. Stereocaulon condensatum Hoffm. 

S. condensatum Hoffm. Deutsch. Fl. 2:130. 1795. 

Primary thallus persistent, formed of coarse, 
intermixed with conspicuous black masses of Stig 
are formed dark, rugose cephaJodia; podetia 
fastigiately branched above; squamules granular, rarely coralline, 
apothecia well developed: spores 20-28X2.5-3/*, average 23X 
2.6 m-, with pointed ends. — Growing on soil. 

rounded granules, 

— — / 

tout, simple or 


Euxope: Fries Lich. Suec. no. 88; Schaer. Lich. Helv. no. 5 9'> 


Lich. Ital. Suppl. no. 29; Nyl. and Norrl. Herb. Lich. Fenn. no. 87 

Canada: Labrador, A. P. Low July 21, 1896 (Can); Quebec, Macon 
Aug. 11, 1905 (Can). 


Massachusetts: Wellesley, Clara E. Cummlngs in Dec. N. A. L. 26; 
South Sudbury, C. M. Carr (R); Ipswich, Wm. Oakes (Tuck); Duxbury, 

/. L. Russell (Tuck); New Bedford, H. Willey (BSNH); Mattapoisett, H. 

Willey (R) . 

Connecticut: Montville, W. A. Setchell (Tuck); Norwich, W. A. Setchell 

The differences which distinguish this species from S. pileatum are suffi- 
ciently indicated in the notes under the latter, to which reference may be made. 

Sect 'on Eustereocaulon 

Primary thallus soon disappearing f podetia mostly tall, well 
developed, and irregularly branched; squamules well developed, 
cartilaginous, gray or w T hite, but never chalky. — Spores 15-35X 
2 • 5~5 P, with some minor variations among the species, but on 
the whole so uniform as to be without value in specific diagnosis. 


S. coralloides Fries Lich. Suec. no. 118. 1817; and Sched. Crit. p. 24. 1824. 
S. corallinum Laurer in Fries Lich. Eu, p. 201. 1831. 
S. corallinum Fries in Tuck. Syn. Lich. N.E. p. 45. 1848. 
S. Despreaultii Delise ms. in Nyl. Syn. p. 249. 1858 (according to authentic 
specimen in Herb. Tuck.). 

Podetia solitary or caespitose, erect, branched, usually exten- 
sively, glabrous or subtomentose; squamules coralline and branch- 
ing, giving the plant a fibrillose appearance; apothecium medium 
to large (1-4 mm.), mostly terminal;] cephalodia of the type of 
S. paschale (q.v.), usually abundant and conspicuous. — Growing 
on rocks (fig. 5). 


Europe: Fries Lich. Suec. no. 118; Stenh. Lich. Suec. no. 82; Sweden, 
Tk. Fries, Leigh ton Collection (NY), also Nylander (loc. cit.); Schaer. Lich. 
Helv. no. 261; Hepp Lich. Eu. no. 114; Pyrenees, R. Spruce in Leighton 
Collection (NY). 

British America: Labrador, Toumsend July 1906 (H); Newfoundland, 
Waghorne no. 46, no. 66 as 5. tomentosum, no. 67 as 5. paschale (CEC) ; New 
Brunswick, Farlow July 1892 (H); Willey (Tuck); Cape Breton Is., Macoun 
July 22, 1898 (Can); Nova Scotia, A. H. Mackay (BSNH); Ontario, Macoun 
(Can); Lake Superior, Macoun (Can); Vancouver Is., Macoun (Can); Colder 
(BSNH); Lya//(Tuck). 

Alaska: Unalaska, /. M. Macoun June 4, 1897 (Can); St. Paul Is., 
J - M. Macoun June ig, 1897 (Can). 



Maine: Eastport, Farlow (H); West Pembroke, Maude C. Wiegand 
Aug. ii, 1909 (W); Mt. Katahdin, J. F. Collins in Herb.; Mt. Desert Is., 
M. L. Wilson (Mt. D). 

New Hampshire: White Mts., Tuckerman Lich. Exs. no. 94; E. Faxon, 

as 5. denudatum (CEC); Clara E. Cummings (R); Farlow (H). 

Vermont: Brattleboro, C. C. Frost (UVM). 

North Carolina: Buckley (NY); /: W. Harshberger (NY). 

A beautiful species, entirely distinct from all others of the boreal group in 
the coralline or fibrillose squamules (fig. 8) , clearly separating it from S. paschale 



of the three species, it is nevertheless rather remarkable that no specimens have 
been seen from the central or western parts of the United States. 

4. Stereocaulon paschale (L.) Ach. 

Lichen paschalis Linnaeus Sp. PL 2 : 1 1 53. 1753 (cf. Wainio Revisio lichenion 
in herbario Linneai asservatorum in Medd. Soc. F. et Fl. Fenn. 14:1. 1886). 

Stereocaulon paschale Acharius Meth. Lich. p. 315. 1803, in part (cf. 
Th. Fries Monog. Ster. p. 58). 

Stereocaulon paschale Fries emend, in Lich. Eu. p. 202. 183 1. 




tomentose; squamules palmate- 


never coralline; apothecia o. 5 to 4 mm. in diameter, average 1 mm., 


gray, spherical, rugose-plicate, containing Stigonema, sometimes 
fibrillose and black. — Growing on rocks, rarely on the earth. 

specimens examined 


man 1842 (Tuck). 

British America: Labrador, A. P. Low July 1896 (Can); M agnorn*, 
Aug. 1891 (NY); Quebec, Gaspe County, /. F. Collins July 19, 19 06 (* ); 

rick, G. U. Hay July 1884 (Can); Nova Scotia, Mackay (BSXHj; 
Ontario, many localities, Macoun (Can); Lake Superior, Macoun Canad. 
Lich. no. 45; Alberta, H. L. Bolley Aug. 1901 (F); British Columbia, Macoun 

New Bruns 

(Can) . 

Alaska: /. .1/ 
Cooley Aug. 5, 1891, as 



1899, Harriman Exped. no. 1299, as 5. tomentosum (CEC); Tort Cla en 


Behring Strait, Trelease July 12, 1899, Harriman Exped. no. 1266, as S. tomen- 

tosum (CEC). 

Maine: Eastnort W 

. Farlow 1877 (Tuck); Portage, L. W. 
W. Riddle Aug. 1004 ftO: Orono. L. W 

July 1907 (R) ; Mt. Desert, M . L. Wilson (Mt. D), Sam Surrey (R) ; Rockport, 
G. K. Merrill Lich. Exs. no. 40; Cumberland, /. Blake (NY). 

New Hampshire: White Mts., Tuckerman Lich. Exs. no. 122; North 
Woodstock, Clara E. Cummings in N. A. L. no. 25, L. B. A. no. 151 ; Crawfords, 
Miss Minns (W); Warren, Charles W. Riddle Aug. 1908 (R); Mt. Chocorua, 
H. H. Bartlett Sept. 1906 (R); Mt. Monadnock, W. G. Farlow (H); R. H. 
Howe, Jr. (Th); Fitzwilliam, R. H. Howe, Jr., Lich. Nov. Ang. no. 22. 

Vermont: Mt. Mansfield, C. G. Pringle 1879 (UVM); Willoughby, W. G. 
Farlow (H); Bristol, L. W. Riddle Aug. 12, 1908 (R). 

Massachusetts: Pepperell, L. W. Riddle May 30, 1909 (R); Mt. Watatic 
and Concord, R. H. Howe, Jr. (Th) ; Arlington, B. Fink May 1895 (R) ; "near 

ton," T. P. Adams (BSNK) 
Connecticut: Woodbridge, 

Mrs. C, W. Harris 1000 (R); L. W 

Aug. 1908 (R). 

Ohio: Lesquereux? (NY). 

Michigan: E. T. Harper Aug. 1899 (R). 

Wisconsin: C. F. Baker (CEC). 

Minnesota: Macmillan 1894 (R); B. Fink July 30, 1902 (R). 

Oregon: E. Hall 1871 (Tuck). 

Tuckerman, following Th. Fries, cites E. Fries as the author of this 
species. The combination was first made, however, by Acharius {op. cit.) y 
who refers to the figure in English botany, pi. 282. According to the Vienna 
Code (Articles 40 and 41), Acharius should be cited therefore as the author, 
even though his conception of the species was later emended by Fries through 
the segregation of S. tomentosum and S. coralloides. 

S. paschale is the commonest and most widely distributed species of the 
genus, at least in the eastern portion of North America. In the west it seems 
to be comparatively rare, its place being taken by S. tomentosum. 

The two species named may be best separated from their allies by the 




tomentosum. Typical S. paschale is said to have glabrous podetia, while 


But a study of a suffi- 

cient amount of material shows that there are all gradations from wholly 

medium amoun 

As the tomentum is the chief character hitherto used as diagnostic 


ith such variability much confusion must result. A second character 
has beeit used is the Dosition of the aoothecia, in typical 5. paschale 




the apothecia being at least in part terminal, while in S. tomentosum they are 
said to be always lateral. Here again we have a variable character which 

cannot be used as a precise criterion. 

In searching for a character which would by its nature be mutually exclusive 
for the two species, the attempt was made to use the cephalodia. The differ- 
ences between the cephalodia typical of these two species are sufficiently indi- 
cated in the descriptions, to which reference may be made. To establish the 
taxonomic value of this structure, it was necessary to prove three points: 
first, both types of cephalodia must not occur on the same plant; secondly, 
the cephalodia must be constant enough in occurrence to be of use ; and thirdly, 
the characteristics of the cephalodia must be correlated with the other diag- 
nostic characters. 

As has been shown by various investigators, the cephalodia of lichens are 
peculiar gall-like outgrowths formed by the hyphae of the thallus surrounding 
some foreign alga. A priori we should hardly expect that such a structure 
would be constant enough to be used in classification. The method adopted 
to prove the points mentioned was to make a statistical study of a large number 
of specimens of the paschale-tomentosum group, noting the type of cephalodium, 
the amount of tomentum, and the position of the apothecia, and then to tabu- 
late the results. In this statistical study material was examined from various 
parts of North America and Europe, and the specimens were taken as they 
came in the herbaria, in order to avoid influencing the results by conscious 
choice. The only exception was made in the case of a few specimens wnicn 
were so clearly depauperate that they could in no way be looked upon as aid- 
ing in the object sought. The results are given in the following tables: 

Number of specimens examined . 
Specimens with both types of cephalodia 
Specimens lacking cephalodia . 
Specimens lacking apothecia . 





of cephalodia 

found shows that for practical purposes the two types are mutually exclusive. 
Cephalodia being present in a larger proportion of specimens than were apothe- 
cia, it follows that if apothecial characters or spore characters are of value 
cephalodia can equally well be used, as far as constancy of occurrence is con- 
cerned. To show correlation of characters, the results may be arranged as 

follows : 






Tomentosum type .... 
Paschale type 



3 ' 

ig io] 


2 95 







Tomentosum type .... 
Paschale type 





These tables show conclusively the extent to which S. paschale and 5\ 
tomentosum intergrade. They show also the confusion introduced by taking 
the presence of tomentum alone as a criterion, for, as is shown in the second 
table, out of 93 specimens 35 were intermediate in the amount of tomentum. 
With such intergrading, complete correlation of characters is not to be expected. 
But the tables show that the types of cephalodia are correlated to a marked 
degree with the two characters that have been recognized as of chief importance 
in separating these two species. The results of the statistical study, therefore, 
support very strongly the suggestion that the cephalodia, when present, are 
of great taxonomic value. Yet taking all available characters into considera- 
tion, there are still specimens which cannot be placed absolutely under either 

characters of both. And this must necessarily 



evidently form a continuous, variable series, the forms that we recognize as 





This is 

40- Stereocaulon paschale var. conglomeratum Fries. 

Fries Sched. Crit. ad Lich. Suec. 3 : 2o. 1824. 

Podetia much branched, forming compact cushions which are 
closely adnate to the substratum, squamules more granular and 
crowded than in the type; usually sterile.— Growing on rocks. 


Europe: Fries Lich. Suec. no. 89; Hepp Lich. Eu. no. 304; Sweden, Are- 
schoug (Tuck). 

New Hampshire: Mt. Monadnock, W. G. Farloiv July 1, 1896 (H); 
Alton Bay, C. /. Sprague (BSNH) . 

Vermont: Mt. Ascutney, R. H. Howe, Jr., Aug. 25, 1909 (Th). 

Massachusetts: Annisquam, Clara E. Cummings 1892 (CEC); Wellesley, 
Clara E. Cummings Dec. 1884 (CEC); L. W. Riddle Jan. 1908 (R). 

This is a reduced form of S. paschale, occurring either at low altitudes or 
on rocks with extreme exposure. It passes into the laxer, more typical form 
by insensible gradations. The only other species with which this variety may 




be confused is S. denudatum, and the specimens from Ipswich, Mass. (Wm. 
Oakes), and from Nantasket, Mass. (H. Willey), in the Tuckerman Herbarium, 
labeled denudatutn, seem to be much nearer this variety of S. paschale. 
The two have a general resemblance in habit, but 6*. paschale var. conglomera- 
tum is less denuded and always has indications of the palmate-digitate type 
of squamules typical of S. paschale. 

5. Stereocaulon tomentosum: Fries. 

S. tomentosum Fries Sched. Crit. ad Lich. Suec. 3:20. 1824. 

Figs, i, 2, 3. 


i, collected by W. Trelease 
um of Clara E. Cummingj 

899, Harriman 

fig- *> 

n. col- 

nat. size 

Uon tomentosum Fr., large form; specimen from Cascade Mts., \ 

7 C. G. Pringle, Sept. 20, 1881, in herbarium of Wellesley College; pho • 

; fig. 3, Stereocaulon Wrightii Tuck.; specimen from St. Michael, ^ Ia !^ 





Podetia loosely attached to the substratum, erect, or spreading, 
3 to 9 cm. high, average 5 cm., more or less densely tomentose; 
squamules palmate-digitate, abundant, more or less crowded; 
apothecia typically minute, o. 5 to 1 mm. in diameter, rarely larger, 
reaching 2 mm., lateral; cephalodia minute, verdigris-green, con- 
taining Nostoc. — Growing on the ground or among mosses, rarely 
on rocks (fig. 1). 




1854 (Tuck); Sweden, Nylander 
k America: Labrador, Waghor 
Macoim Auer. 22. 1883 (Can): "P 


1882 (Can); Gaspe County, Quebec, Macoim July 28, 1882 (Can); /. F. 
Collins Aug. 17, 1906 (R); Cape Breton Is., Macoun July 9, 1898 (Can); 

Macoun J 





June 13, 1007 (Can) ; L. R. Waldron J 

Alaska: Dr. Kellogg (Tuck); P. M. Newhall on Univ. Calif. Exped. 1899 
(CEC); Trelease in Harriman Exped. nos. 1267, 1297, 1300, 1301; Coville 
and Kearney in Harriman Exped. nos. 609, 981, 21 21 (CEC). 

Maine: Eastport, Farlow (H); Mt. Desert, M. L, Fernald Sept. 18, 1892 
(CEC); Cumberland, J. Blake Dec. 1855 (CEC). 

New Hampshire: White Mts., Tuckerman Lich. Exs. no. 23; Crawfords, 
Miss Minns (W); North Woodstock, Clara E. Cummings Aug. 19, 1892 (CF.C); 
Shelburne, Farlow Aug. 1894 (H) ; Fitzwilliam, R. H. Howe, Jr. (Th.) 

Vermont: Hinesburgh, C. G. Pr ingle 1878 (UVM); Brattleboro, C. C. 
Frost (UVM). 

Massachusetts: Magnolia, C. /. Sprague, as S. paschale (BSNH); 
Duxbury, J. L. Russell (B); New Bedford, H. Willey (B). 

Connecticut: Ellsworth, H. A. Green Aug. 7, 1887 (NY). 

New York: C. F. Austin (NY); E. A. Burt (F); K. M. Wiegand (W); 
L. W. Riddle (R). 

Illinois: /. Wolf 1888 (NY). 

Minnesota: Various localities, B. Fink, in Herb.; Macmillan (F). 

Colorado: T. S. Brandegee (R); F. E. Clements, as S. paschale (H). 


W. Calkins no. 340. as S. paschale (W) 

June 30, 1906 (F) ; Longmire's Springs, E. T. Harper Aug. 1906 


fddt 1882 (Can). 

(H) ; J. W. Eck- 



The relationship of S. tomentosum to S. paschale is sufficiently discussed 
under the latter species, to which reference may be made. The palmate- 
digitate type of squamules will serve to distinguish these species from others 
having the same habit, including 5. alpinum, our only other species with typi- 
cally tomentose podetia. 

Certain material has been examined from our northwest coast, especially 
from the Cascade Mts., Oregon, which seemed at first to be distinct from 
S. tomentosum (fig. 2). These specimens are unusually large and finely 
developed, with large apothecia, and bearing conspicuous white structures 
which at first appeared to be a new type of cephalodium. Examination 01 
these structures failed to show any type of alga other than the Cystococcus 
which forms the normal gonidia for the genus, and the structures cannot 
therefore be considered cephalodia. After careful study I am unable to look 
upon this Oregon material as anything more than exceptionally luxuriant 
S. tomentosum. 

It is otherwise, however, with another form of S. tomentosum, which is 
undoubtedly of varietal rank and may be designated 





mulae paucae et dispersae; tomentum 
in forma typica. — Habitat in k 

simple or nearly so ; squamule 

and scattered; tomentum 
type. — Growing in bare sai 

Type specimens: "On sandy river bottom, Mt. Rainier region, Washing- 





Oregon: Cascade Mts., Moses Craig Aug. 1898 (CEC), A. S. Foster 





1887 (Can). 

This variety is very striking and distinct, and can scarcely be con 
with any other North American form. 

6. Stereocatjlon alpinum Laurer. 

5. alpinum Laurer in Fries Lich. Eu. p. 204. 183 1. ^ and 

S. tomentosum var. alpinum (Laur.) Th. Fr. Comm. Ster. p. 3°- l857 ' 
American authors. 









Figs. 4, 5, 6.— Fig. 4, Stereocanlon tomentosum var. w»/>/ex Riddle; type specimens 
from Mt. Rainier region, Washington, collected by T. C. Frye, Aug. 14, ia°4, in 

L 1 1 t If. ?J T7 1 - 

xi um 

specimen from the Austrian Tyrol, collected by Kernstock and issued in Krypto- 
gammae Exsiccati (Vienna), no. 355, herbarium of L. W. Riddle; photo, nat. size; 
fig. 6, Skreocaulon denudatum Floerke; specimen with the granulate type of squamules, 
from Minnesota, collected by Bruce Fink, June 19, 1897, in herbarium of L. W. 
Riddle; photo, nat. 



Podetia stout, erect, i to 6 cm. tall more or less densely 
branched, and tomentose; squamules in the form of coarse, crowded, 
conglomerate granules; apothecia (rare) medium to large, lateral 
or terminal; cephalodia of the type of S. tomentosum (q.vX — Grow- 
ing on exposed soil, especially in arctic-alpine regions. 


Europe: Authentic specimens ex Herb. Laurer (F); Schaer. Lich. Helv. 
no. 263; Massalongo Lich. Ital. no. n; AnziLich. Ital. Suppl. no. 26; Rabenh. 
Lich. Eu. no. 859; Arnold Lich. Exs. 5 no. 641a; Finmark, Th. Fries (Tuck). 

British America: Greenland, Fink (NY); Cumberland Is., Howgate 
Exped. 1877 (Tuck); Diggs Is. Husdon Bay, R. Bell (Can); Labrador, Wag- 
home no. 20, also no. 21 as S. condensatum (CEC); A. P. Low (Can); Alberta, 
Macoun June 28, 1905 (Can). 

Alaska: St. Paul Is., T. C. Kincaid (F); Is. in Behring Strait, C. Wright 
(Tuck); Port Foulke, Dr. Hayes (Tuck); Fred Funston 1894 (CEC); Trelease 
in Harriman Exped. nos. 1268, 1270, 12980 (CEC); Ball in Harriman Exped. 
no. 1295, a s S. tomentosum (CEC). 

Maine: Eastport, Farlow (H); Portage L. W. Riddle Aug. 1907 ( R )- 

New Hampshire: Mt. Washington, Farlow Sept. 1894 (H). 

Minnesota: B. Fink no. 93 (F). 

Oregon: Mt. Hood, E. Hall 1871 (Tuck). > 

Washington: Mt. Rainier, C. V. Piper 1895 (CEC); Friday Harbor, 
B. Fink June 28, 1906 (F). 

This is clearly distinct as a species from S. tomentosum, as the type 
squamules is a specific criterion throughout the genus. It is most liable to 
confusion with S. denudatum which it resembles in general habit and range, 
but it can be distinguished by the tomentum and the type of cephalodium. 

7. Streocaulon denudatum Floerke. 

. 5. denudatum Flke. Deutsch. Lich. 4:13. 1819. 

original specimen). 


Journ. Nat. Hist. 3:302. 1840 (according to 

r „, solitary or aggregated, shrubby-erect, 

subsimple below, branching above, wholly glabrous, denuded 
below, squamulose above; squamules crowded, in the typica 
form umbilicate, attached by the center, and with the margin 
crenate (see fig. q), or often in the form of coarse, rounded, con- 
glomerate granules; apothecia rare in North American 

rocks or soil (fig. 8). 


Growing on exposed 






Europe: Authentic specimen ex Herb. Floerke in Herb. Tuck.; Fries 
Lich. Suec. no. 346; Arnold Lich. Exs. no. 1576; Sweden, Forssell 1880 (NY); 
Scotland, Leighton July 1869 (NY); 
France, L. Breviere (R) . 

British America: Greenland, Eber- 

lin 1885 (Can); Warming and Holm 1884 
(H); Kikkeston Is., Howgatc Exped. 1877 
(Tuck); Hudson's Strait, R. Bell 1884 
(Can); Labrador, Wm. Palmer 1887 
(NY); Owen Bryant Aug. 22, 1908 (H); 
Newfoundland, no collector given 
(BSNH) ; Delise (Tuck) not typical. 

New Hampshire: White Mts., Tuck- 
erman Lich. Exs. no. 114; Mt. Washing- 
ton, W. G. Farlow Sept. 1891 (H); Mt. 
Adams, W. G. Farlow Sept. 1897 (H); 
Mt. Willard, E. Faxon June 1882 (CEC); 

Mt. Monadnock, W. G. Farlow July 1896 

Vermont: C. C. Frost (BSNH); Mt. 
Mansfield, W. G. Farlow Sept. 1890 (H). 

New York: Mrs., C. W. Harris June 
1000 (R); Adirondack Mts., W. G. Farlow 
Sept. 1902 (H). 



Figs. 7, 8, 9.— Fig. 7, palmate- 
digitate type of squamules found in 
S. paschale and S. tomentosum; en- 

Massachusetts : Mt. Holyoke, TTO- i arg ed; fig. 8, typical squamules 

man May 20, 1839 (Tuck). 

of 5. coralloides; enlarged; fig. 9, 

Minnesota: B. Fink no. 49 in Herb, squamules of S. deuudatum; right, 

Washington : Suksdorf (B SNH) . 

umbilicate type; left, granular type; 

California : Bolander in Herb. Tuck. enlarged and semi-diagrammatic. 


Alaska: Dr. Kellogg 1867 (Tuck); W. A. Setchell, Univ. Calif. Exped. 
1809 (CEC); W. H. Evans 1897, as S. alpinum (CEC); Harriman Exped. 
no- 1279, as S. alpinum (CEC). 

The two types of squamules described for this species intergrade, and the 
granulate type is scarcely worthy of varietal rank. The typical umbilicate 
squamules are found in specimens from arctic or subarctic stations. When 
these are present there should be no difficulty in distinguishing S. denudatum 
from all other species. But specimens with the coarsely granular squamules 
are less distinctive, and are frequently mistaken for either S. paschale or S. 
alpinum. From the latter the wholly glabrous podetia and the type of cepha- 
lodia should serve to distinguish S. denudatum; while the error of mistaking it 
for 5. paschale may be avoided if it is borne in mind that in true 5. paschale 



some indication of the characteristic palmate-digitate type of squamules is 
always to be seen, and this type is never to be found in S. denudatum. 

The most puzzling specimens that I have seen are those collected in New 
Jersey by C F. Austin. Portions of this material have been examined in the 
Tuckerman Herbarium, the Sprague Collection at the Boston Society of Natural 
History, and the Herbarium of the New York Botanical Gardens. It was 
determined by Tuckerman as S. denudatum, and is cited in his Synopsis of 
North American Lichens, p. 233; yet it is scarcely typical of this species, and 
it seems much nearer to S. paschale, a view which would be supported by the 
latitude of the station, 5. denudatum, being essentially an arctic-alpine species. 

According to the description of this species given by Th. Fries, the apo- 
thecia are minute (i.e. about 0.5 mm. in diameter) and lateral. But the speci- 
men collected, by C. C. Frost in Vermont and now in the Sprague Collection 
(BSNH), agreeing in all other particulars with S. denudatum, has terminal 
apothecia, over 1 . 5 mm. in diameter. 

8. Stereocaulon Wrightii Tuck. 

S. Wrightii Tuck. Suppl. II to Enum. N. Am. Lich.. in Am. Journ. Sci. 

28:202. 1859. 

Stereocladium Wrightii Nyl. Lich. Freti Behr. p. 85. 1888. 

907 (see below). 

Stereocaulon foliiforme 




wholly glabrous, denuded below, ending in broad, foliose, convolute 
tips, taking the place of the usual squamules; or -the podetia 
reduced and the expanded portions alone present; apothecia 
unknown; cephalodia doubtful (fig. 3). 

Hitherto this species 

Charles Wright 


Strait, during the U.S. North Pacific Exploring Expedition (i853- l8 5 6 )- 
To this may now be added two other localities, from both of which specimens 
have been distributed under other names. A comparison of these specimens, 
however, with the type specimens of S. Wrightii in the Tuckerman Herbarium 
shows that they are undoubtedly this species. These specimens are from 


J .r J . it ..^vi, i TO3&o, wnccicu uy vv. /\. ^JSiuJiiii,!- 011 uic »""«- -v , 

fornia Expedition in 1899, and distributed as Stereocaulon denudatum; an 
from Shiribeshi, Japan, collected by Abbe Fauiue, July 1905, and 1SSU lD 
Faurie Lich. Jap. no. 6999 as Stereocaulon foliiforme Hue, n. sp.; an authentic 
specimen determined by Hue himself is in the herbarium of W. G. Farlow. 

Section Chondrocaulon {Lcprocaidon Nyl.) 
Primary thallus present or disappearing; podetia reduce.- 
simple or branched; squamules granular, dissolving into chalky- 
white powder. 


9. Stereocaulon albicans Th. Fr. 

S. albicans Th. Fr. Comm. Ster. p. 36. 1857. 


mens) . 

Podetia short, mostly under 1 cm., slender, caespitose, branched, 
crowded and intertangled; squamules minutely granular, becom- 
ing powdery; apothecia unknown. — The whole plant is character- 
istically chalky-white and powdery. 


.North America: Greenland, Kikkeston Is., Howgate Exped. 1877 (Tuck); 
Vancouver Is., Macoun May 19, 1893 (Can); Colorado, T. S. Brandegee 
(BSNH); Arizona, Rincon Mts. near Tucson, /. C. Blunter Oct. 1909 (R); 
California, C. R. Orcutt (BSNH) ; W. G. Farlow (H) ; Guadeloupe Is., E. Palmer 
1875 (Tuck); Cuba, C. Wright (Tuck). 

South America: New Granada, Lindig no. 2502 (Tuck); Peru, Wilkes 
Exped. (Tuck), type of S. tenellum Tuck. 

This is a rare and interesting species, first described by Th. Fries from 
material collected by Gaudichaud in Peru, but with a wide distribution as 
indicated. It is the representative in the western hemisphere of the Old World 
5, nanum Ach. It does not resemble any other American species sufficiently 
to cause confusion. 


Stereocaulon nanodes Tuck, in Suppl. II to Enum. N. A. Lichens, 

1- Jour. Sci. 28:201. 1850. 

Primary thallus absent; podetia about 1 cm. tall, dendroid- 



of small rounded granules, dissolving into fine, whitish powder 
(but not chalky as in S. nanum); apothecia terminal, or absent 
and the podetia ending in masses of soredia as S. pileatum— -Spores 

24-42X2.5-3 (JL. 

I have examined at the Tuckerman Herbarium the type specimens of this 




dily referable to any other sDecies of Stereocaulon 

as doubtful on account of the fact that it has apparently never been collected 


re gions, botanically, in North America. 


i'oundland on the authority of Dr. J. W. Eckfeldt. Through the 



have had an opportunity to examine the specimens upon which this record was 
based. One sheet is S. p ilea turn Ach.; the other, depauperate specimens of 
Cladonia decorticata (Flke.) Spreng. The species is known, therefore, only from 
the type collections. 

Stereocaulon ramulosum (Sw.) Ach. 

In the herbarium of the Geological Survey of Canada is a sheet bearing 
three specimens of this species with the label: "Lichenes Boreali Americani. 
No. 89. Dalles, Oregon, on the earth. Legi John W. Eckfeldt. 1880 
As no corresponding specimens are to be found in Dr. Eckeeldt's herbarium 
at the Philadelphia Academy of Sciences, and as S. ramulosum is not otherwise 
known from the United States, there has undoubtedly been some mistake in 
the labeling of the specimens mentioned above. 

Wellesley College 
Wellesley, Mass. 





(with two figures) 

The genus Frenelopsis was founded by Schenck in 1869 1 upon 
abundant material from the Wernsdorferschichten (Lower Cretaceous), 
and named from its resemblance to the modern genus Frenela. Thuites 
Hoheneggeri of Ettingshausen was the type and only species. This was, 
he says, the most abundant fossil in those beds in which it occurs, and 
it received a careful and elaborate treatment at his hands. This species 
has subsequently been recognized in the Kome beds of Greenland, the 



In 1880 Hosius 



described an additional species, F. occidentalism from Portugal, which 
S aporta has shown 4 ranges from the Urgonian of Cereal, through the 
Albian of Nazareth, into the Cenomanian of Alcantara and Padrao. 
The latter author also describes an additional species, F. leptoclada, 5 
which is confined to the Lower Cretaceous of Portugal (Neocomian- 

In 1889 Velenovsky described the Bohemian form F. bohemica from 
the Cenomanian of that country {op. cit.), and the next year Fontaine 
described the two species F. ramosissima and F. parceramosa from the 
Potomac group of Virginia, the same author three years later founding 
a third species, F. varians, upon material from the Trinity group of 
Texas. Newberry in 1896 described a ninth species, F. gracilis, 6 

1 Palaeontographica 19:13. 1869. 

2 Ibid. 26:132, 181. pi. 37, fig. 148. 1880. 

3 Cont. Foss. Fl. Port. 188 1 : 21. pi. 12, figs. 3b, 4-7. 

4 Fl. Foss. Port. 1894: 139, I99 , 214. pi. 26, fig. 16; pi. 36, figs, r, 2; pi. 38, figs. 

**j; pi. 39, fig. 20. 


5 Ibid. 109, 113. pi. i 9 ,fig. 18; pi. 21, figs. 9-1 1. 

6 Fl. Amboy Clays 1896:59. pi. 12, figs. 1-311. 

[Botanical Gazette, vol. 50 


which is a very abundant Upper Cretaceous type, and which has been 
recently shown by Hollick and Jeffrey to be unrelated to FrenelopsisJ 
Although fruiting specimens have not been found, the position of 
the genus in the Cupressineae is not disputed at the present time, 
although at one time Heer argued for an affinity with Ephedra. The 
genus may be defined as follows: 

Shrubs or trees with cylindrical, jointed, monopodial stems and branches, 
the latter of which may be alternate, apparently in a single plane, or whorl, 
often of large size, stems up to 5 cm. in diameter having been found in the 
Virginia area. Leaves much reduced, somewhat variable in outline, in general 
triangular with a broad base and an acute apex, squamiform, appressed, one 
to four at the nodes, decussate. Internodes variable in length, but 
longer in the apparently annual shoots, which were more or less deciduous 
and functioned as leaves, since the fine longitudinal striae with which their 
surface is ornamented turn out to be rows of stomata in certain of the species 
which have been examined microscopically. . 

It is for the purpose of describing the latter characters of this inter- 
esting American Low r er Cretaceous species that the following brief note 
is published. 



far confined. It is by far the most abundant form at the celebrated 
plant locality of Fredericksburg, Va. The coarse arkosic sandy clays of 
this age are sometimes packed with the remains of this species, with its 
crowded twigs and short internodes generally completely flattened, and 
w T ith all of the tissues gone except the epidermis, which must have been 
very tough and coriaceous in life, since the preservation of these forms 
was largely due to its resistant nature. The cuticle of the type of the 
genus, F. Hoheneggeri, was studied by Zeiller and described in 1882, 8 and 
six years later Velenovsky described^ the epidermal features of F. 
bohemica. The stomata in these species were found to consist of usually 
four cells, although sometimes five or even six were present. These 
guard cells are symmetrically arranged, the opening between them being 
approximately in the form of a narrow-rayed star. According to the 
former author, they ally these forms with the existing species of Cal- 

7 Mem. N.Y. Bot. Gard. 23:6. 1909. 

8 Obs. sur quelques cuticules fossiles. Ann. Sci. Nat. Bot. VI. 13:231. P l - JI ' 


9 Ueber einige neue Pflanzenformen der bohmischen Kreideformation. Si z- 
K. Bohm. Gesell. Wiss. Prag 1888:590. figs. 1-3, 10. 




litris and Libocedrus, and effectually disprove Heer's contention that 
this curious genus is a member of the Gnetales allied to Ephedra. 

In F. ramosissima we find a very similar arrangement to that described 


in the two species just mentioned. The epidermal cells are very small, 
the largest not exceeding 0.025 mm. in diameter, and the average being 
about half this size. They are roughly rectangular in shape and have 
very thick walls. Their most curious feature, one not observed in any 




Fig. i 




-»«uvvj YO.LJ ill ^JUJllllllCll^V, 11VA111 UiUin, ^jv^m.- ~- ^ 

to pointed spines o . 025 mm. in length. These are not present on all 
°f the epidermal cells, and some preparations of the epidermis are 
apparently entirely smooth. Fig. 1 shows a characteristic bit of the 
epidermis dotted with these spines. For the camera lucida drawings 




of the spines, probably all of them, have a central cavity opening into 
the interior of the epidermal cell, which they surmount, as is shown in 

3 o8 



one of the individual spines figured. The second and third single spines 
figured show irregular cavities toward the apex which are apparently 
cut off from the cell cavity, and the third spine figured gives a good idea 
of the papillose character of those adjacent cells in which these processes 
are not prominently developed. 

In the area included in fig. i are three of the curious stoma tal open- 
ings which apparently characterize the genus Frenelopsis. These are 
circular in outline and about 0.03 mm. in diameter. They are very 
numerous, but whether they are localized on certain portions of the 
branches which perform the functions of leaves in this genus, or whether 
they are uniformly distributed on the annual shoots, could not be deter- 
mined. They consist of five or six guard cells arranged around the 

Fig. 2 

central stomatal opening. These cells are much thinner-walled than 
the epidermal cells. In form they are relatively slender distally and 
broad proximally. As viewed through the microscope, they are darker 
colored around the stomatal opening and peripherally they are lighter. 
Since structural material is not available, their exact attitude is made out 
with difficulty. Their outer centrally directed ends come into focus 
at about the same time as do the outer ends of the longer spinelike pro- 
cesses, or very soon after, while their inner broad ends are visible after 
the epidermal cells have gone out of focus ; hence it is obvious that they 
are inclined toward each other and project outward for a considerable 
distance from beneath the surrounding epidermal cells. It is believed 
that fig. 2, which is a diagrammatical drawing of a single group of these 
guard cells and two adjacent epidermal cells viewed in a radial section 
of a twig, gives an accurate idea of their arrangement and proportions. 




In their more essential characters they agree with the stomata as described 
by Zeiller for F. Hoheneggeri and by Velenovsky for F. bohemica. Just 
what were the physiological factors responsible for the great reduction 
of the leaves and the assumption of the photosynthetic processes by the 
branches in Frenelopsis it is difficult to imagine. Such features are 
usually associated with peculiarities of climate and habitat, and suggest 
strong insolation and lack of humidity; but such conditions are not 
suggested by the other members of the flora associated with Frenelopsis, 
since with the Potomac species are found large numbers of ferns, many 
of them apparently tree ferns with decompound fronds a meter or more 
across, and large numbers of cycads of various genera and large size; 
while in the latest beds in which F. ramosissima occurs there are con- 
siderable numbers of dicotyledonous leaves, some of which are allied 
with genera which in the modern flora are confined to tropical areas 
where the humidity is high and the rainfall heavy. 

It is possible that these peculiar features in the cretaceous species of 
Frenelopsis were inherited from triassic ancestors which acquired them 
during those portions of the Triassic when the climate was extremely 
arid, as we know it was from physical as well as paleontological criteria. 

Edward Wilber Berry, The Johns Hopkins University, Baltimore, 



The teaching botanist 

The second and revised edition of this book 1 is almost completely rewritten 
and is larger than the former edition by more than 60 per cent. The general 
plan is quite similar to that of the former book, which was reviewed in this 
journal eleven years ago (28: 276. 1899). During the intervening years atten- 
tion to the teaching of science has greatly increased; and the author has been 
a potent factor in stimulating a scientific attitude toward science teaching. 
Although the results of experiments that have been included in this volume 
have not furnished solutions to many of the difficult problems, some progress 
has been made in the direction pointed out in the introduction to the first 
edition, in which the author said: "The botanical course of the near future must 
be more adaptive to education, more broad and representative of the science, 
more economical of energy than in the past." 

The book consists of two parts and an appendix. Part one includes nine 
chapters which deal with the purpose and methods of botanical teaching. It 
treats of such topics as the place of science in education, the kind of botany that 
is of most educational worth, desirable characteristics of good teaching and good 
teachers, the equipment and method of work in the laboratory, botanical 
books, etc. These discussions are of the greatest value to anyone who teaches 
botany, and some of the chapters will be found equally helpful to teachers ot 
other sciences. The discussions are broad, free from a didactic attitude, ana 
full of stimulus to new endeavor on the part of those who are engaged in teach- 
ing science. The practical suggestions regarding laboratory appliances an 
the purpose and means of experimentation are almost indispensable to pro- 
gressive teachers. 

Part two consists of "outlines and directions for a synthetic course in the 
science of botany." It is divided first into the "structure and function of 
plants," a commendable change from the former titles "the principles of the 
science of botany," and "natural history and classification of the groups 
plants." The order of the topics within these divisions is essentially the same 
as in the first edition, but the outlines for work are greatly changed. They 
are more complete and more definite, more easily interpreted by the studen , 
and at the same time better calculated to secure independent investigation 
and inference. Although the outlines purport to represent a synthetic course, 

1 Ganong, William F., The teaching botanist. Revised edition, pp. xi+439- 
jigs. 40. New York: Macmillan. 1910. 




the morphological, ecological, and physiological treatment of each topic is 
kept rather distinct. At times excellent work that is suggested in physiology 
is quite unrelated to the other work with which it appears, as when the physi- 
ology of turgidity and osmosis is interpolated under the caption of " the i 
phology of flowers," or the tropisms under "the morphology of fruits." 

* There is confessedly presented more work than can be done in a year. The 
teacher must adjust the course to his needs and facilities. The outline will 
doubtless prove most valuable in directing many teachers so that they can 
arrange the kind of course that best suits their needs. An appendix gives 
the unit statements for courses in secondary schools as they have been out- 
lined by the special committees, one from the Botanical Society of America 
and the College Entrance Board, and the other from the North Central Asso- 
ciation of Colleges and Secondary Schools. These are the two representative 
organizations that have attempted to outline these statements, and therefore 
their units should be of particular interest to botanists. 

Teachers of botany and botanists in general are greatly indebted to Pro- 
fessor Ganong for this book, which should be influential in enhancing the 
educational efficiency of science and particularly of botanical science. — O. W. 

Tropical agriculture 

Agriculture in the tropics is the title of a book 2 designed to treat chiefly 
of the commercial aspects of tropical plants, and is in no sense a guide to the 
practice of agriculture. The most valuable arable lands of the tropics are now 
under the control of white people, and the book obviously is intended to furnish 
them with data that "may be helpful and thought-stimulating for the student, 
administrator, or traveller." The great influence of "western" civilization 
and agriculture is credited with having brought a revolution in .tropical agri- 

Parts I and II (" The preliminaries to agriculture" and "The principal culti- 
vations of the tropics") will be of interest to students of plant life, while these 
parts and parts III and IV ("Agriculture in the tropics" and "Agricultural 
organization and policy") will interest geographers and economists. Topics of 
special interest in part I are land and soil, climate, drainage and irrigation, and 
plant life in the tropics (acclimatization). In part II there are presented the 
leading tropical economic plants and plant products, their botanical nature, 
history, cultivation, productivity, use, marketability, commercial importance, 
adaptability to new regions, and dangers from plant and animal parasites. In 
part III much attention is given to the need of giving modern agricultural 
education to the peasants who farm the tropical estates. 

The book contains an immense amount of statistical matter relative to the 


2 Willis, J. C, Agriculture in the tropics; an elementary treatise, pp. xvjii-f 
222. fig S , 25. Cambridge Biological Series. London: Cambridge University Press. 


importance of agricultural plants and their future possibilities. It will be 

found a valuable reference book upon many questions pertaining to economic 
and commercial aspects of tropical plants. Botanically, however, the book 
is often defective, as for example, in speaking of the growth of Cannabis saliva 
for its opium-like drug, the author says: "The male flowers are removed in 
November, for if the female flowers are fertilized there is no formation of the 
drug." — O. W. Caldwell. 

The geography of ferns 

It is a praiseworthy thing for an investigator, who has devoted years to 
taxonomic exploration, to bring together in readable form the many things of 
geographic interest which he has observed. It is exactly this service which 
Christ, the well-known student of the ferns, has now performed. 3 The volume 
is divided into two parts, corresponding somewhat to the usual divisions of 
ecological and floristic geography. Christ regards the ferns as controlled 
by the same general distributional factors as the seed plants, the most note- 
worthy difference consisting in the pronounced tendency of ferns to be hygro- 
phytic mesotherms. The great fern areas of the world are essentially coin- 
cident with the forest areas, very few species existing where the rainfall is less 
than 60 cm., and the greatest development occurring where the rainfall is over 
200 cm. 

The edaphic conditions under which ferns live are first noted, attention 
being called to the fact that most species are humus forms, and but slightly 
dependent on the mineral nature of the soil. Under the head of climatic 
conditions, a number of characteristic fern formations are described. The 
hygrophytic ferns are treated at considerable length, especial attention being 
devoted to the epiphytic forms. The features of the xerophytic ferns are well 
portrayed. In the floristic part of the work, consideration is given to a number 
of cosmopolites, and also to endemic forms and to species with disconnected 
areas. The body of the second part is made up of the treatment of the floristic 
regions of the world. Here, as elsewhere in the volume, the author makes it very 
clear that the ferns, in spite of their great age, are far from being a senescent 



all botanists. 


ecological and geographic facts concerning most living ferns— Henry v>. 


An organic chemistry 

The third English edition of Holleman's Organic chemistry has just 
appeared,* edited by A. Jamieson Walker. The value of the book as a text 


3 Christ, H., Die Geographie der Fame. 8vo. pp. 358, with frontispiece, A* 
129 (mostly photographic reproductions), and 3 maps. Jena: Gustav Fischer. *9 


80. New York: John Wiley & Sons. 1910. 


is shown by a statement from the author's preface: "Besides the four editions 
of the original Dutch volume and three in English, seven editions of this book 
have been published in German, two in Russian, two in Italian, and one in 
Polish. A French edition and a Japanese edition are also in preparation.' 
A second quotation from the preface gives the aim of the book, which is well 
carried out, and it gives the text its great value as a general reference book' 
for students of physiology. "Most of the short textbooks of organic chemistry 
contain a great number of isolated facts; the number of compounds described in 
them is so considerable as to confuse the beginner. Moreover, the theoretical 
grounds on which this division of the science is based are often kept in the 
background; for example, the proofs given of the constitutional formulae 
frequently leave much to be desired. However useful these books may be for 
reference, they are often ill-suited for textbooks, as many students have learned 
from their own experience. " 

The chapters on sugars, amino acids, and proteins, of the greatest direct 
interest to physiologists, though brief, are certainly clear statements of the 
fundamental facts of the chemistry of these bodies. The chapter on proteins 
is in the body of the book just after amino acids, instead of in an appendix, 
as it appeared in the second edition. Walker is credited with having intro- 
duced into the book the protein classification adopted by the Chemical Society 
of London, the English Physiological Society, the American Physiological 
Society, and the American Society of Biological Chemistry. — William 


Magazines for students of genetics. — The era of experimental study in 
heredity and evolution has called for new publications devoted to the results 
of research in this field. While all the biological journals occasionally contain 
articles which are of interest to the student of experimental evolution, several 
special magazines have been established which are quite indispensable to any- 
one who wishes to keep reasonably well informed regarding current progress 
in genetics and related subjects. 



for the publication of papers on mathematical methods of dealing with varia- 
tion, heredity, selection, etc., and the results of their application. While 
not many of the articles published in Biometrika deal strictly with genetics, 
it was the first journal to voice the demands for more exact methods of investi- 
gating problems of evolution, and as the whole trend of modern biology is 
toward the greater exactness involved in mathematical treatment of biologi- 
cal data, Biometrika should continue to fill an increasingly important place, 
notwithstanding the unfortunate fact that there is a tendency of late to allow 
Personal feeling: to dominate both the policy of the magazine and the attitude 


of many prominent students of genetics toward it and the methods it advo- 


The second magazine which is devoted specifically to evolution and related 
subjects is the Archiv fur Rassen- und Gesellschafts-Biologie, edited and pub- 
lished in Miinchen by Dr. A. Ploetz, assisted by Dr. L. Plate and several 
others. This magazine was established in 1904, just at the time when interest 
in evolution was being revived by the general introduction of the new methods. 
It is not wholly occupied with the results of experimentation, but contains 
besides these many philosophical and more or less academic discussions, which 
give it the flavor of transition from the older evolutionary literature of the 
post-Darwinian writers to the new type of literature introduced and exempli- 
fied by the w r ork of De Vries and Mendel. 

In 1908 a new monthly German magazine entitled Zeitschrift fur Induk- 
tive Abstammungs- und Vererbungslehre was established in Berlin under the 
editorship of Dr. E. Baur, assisted by Drs. Correns, Hecker, Steinmann, 
and Wettstein. This magazine is at the present time the most valuable 
of all similar journals, as it offers in excellent form a large volume of original 
matter, adequate abstracts from a large proportion of the papers on the same 
subjects published in other journals, and finally gives a very complete classi- 
fied list of new literature dealing with the same subjects. This journal is 
absolutely indispensable to all who wish to investigate problems of heredity, 
variation, and evolution. 

In October 1909 a journal first made its appearance in London entitled 
the Mendel Journal, which is unique in a number of particulars. In the tirst 
place, its editorship is entirely anonymous, though many internal evidences 
leave little doubt as to the source of the dominating influence. It seems to 
be in a sense the organ of the Mendel Society of London, as it publishes the 
various addresses made before that body. While this journal has a number 
of interesting papers which supply valuable data, especially on human inherit- 
ance, its whole attitude is one much to be regretted. The express object of 
the magazine is to furnish a propaganda for Mendelism. It represents the 
"Mendelians" as an army which is fighting an opposing army, the "Biome- 
tricians," and makes many unwarranted slurring attacks upon Professor 
Pearson and those who have assisted him in working out the biometnca 
methods of dealing with biological data. No true scientist can fail to depre- 
cate the introduction of propagandism and personal enmity into science, 
may be expected that the publication of such a journal will act as a boomerang, 
and will do the cause of Mendelian investigation a distinct injury. The aim 
should be simply to arrive at the truth and not to adopt and establish a cree 
to be forced upon others who cannot be convinced of the truth by the da 
presented. Like Biomdrika, the Mendel Journal has no regular periodicity, 
but its appearance is promised whenever sufficient material is at hand. 

In America the demand for journals dealing with genetics has broug 
a most satisfactory response from the American Naturalist, which about wo 


years ago, under the editorship of Dr. J. McK. Cattell, adopted as a general 
policy a specialization upon evolutionary topics. The American Society of 
Naturalists chose for itself the same field, and the American Naturalist is in 
a certain sense the organ of that society. 

The last arrival in the field is the new magazine issued by the American 
Breeders' Association, and known as the American Breeders' Magazine, pub- 
lished in Washington under the editorship of the secretary of the Association, 
W. M. Hays, assisted by the secretaries of the Plant and Animal Sections, 
N. E. Hansen and H. W.. Mumford. The first issue of this magazine has just 
appeared, and gives every evidence of its intention to take a dignified place 
among economic journals, and a modest and unassuming place among scien- 
tific magazines, thus correctly representing the unique position of the Asso.- 
ciation whose organ it is. The American Breeders' Magazine is being issued 
as a quarterly, but it is hoped by those who have it in charge that it may soon 
be changed to a monthly magazine. The opening number presents as a 
frontispiece a portrait of Charles Darwin, and also gives portraits of Gregor 
Mendel and Amos Cruikshank, the latter the originator of the shorthorn 
cattle, and said to be the first to utilize fully the discovery of a mutation 
in the establishing of an important economic breed. Besides brief bio- 
graphical sketches of these three men, this first issue presents papers on 
" Increasing protein or fat in corn," by L. H. Smith; "New methods of plant- 
breeding/' by George W. Oliver; "The army horse," by Carlos Guer- 
rero; "Imperfection of dominance," by C. B. Davenport; "Poultry-breed- 
ing in South Australia," by D. F. Lowrey; and several articles on the breeding 
of deer and other wild animals by D. E. Lance and other members of the 
committee on breeding wild animals. As it is the plan of the American 
Breeders' Association to continue the publication of its Year Book, the issue 
of this magazine will make it even more important that all those who are 
engaged in investigation or who are interested in matters involving genetics 
should become members of the Association and thus secure its publications.— 

Geo. H. Shull. 

Respiration . — Palladin 


similar tissues that have been 
He finds that 0.09 per cent quinine 

hydrogen chlorid more than triples the C0 2 output from living stem tips 

while it does not affect the kill 

less effective on the living tissue and indifferent in the dead; and 1 .0 per cent 
tripled the product from the living and reduced it markedly in the dead tissue. 
In the living: bulbs of Gladiolus Lemonia, 8 cc. of ether per 7.5 l Jters of air 

s Palladin, W., Ueber die Wirkung von Giften auf die Atmung lebender und 
abgetoteter Pflanzen, sowie auf Atmungenzyme. Jahrb. Wiss. Bot. 47:431-461. 


doubled the C0 2 production, while 16 cc. in the same volume was less effective. 
Neither increased the output from killed bulbs. Ether gave little stimula- 
tion in the living bulbs of Gladiolus Colvillii and Allium Cepa, while it cut 
greatly the output from dead tissues. Arbutin (i and 2 per cent) reduced 
considerably the C0 2 yield from live wheat seedlings, and very markedly that 
from killed ones. Palladin finds that the stimulative or inhibitory effect 
of poisons on the living tissues was not accompanied by an increase or decrease 
in peroxidase, but the decreased output from dead tissues was accompanied 
by a decrease in peroxidase. The stimulatory effect of poisons gradually 
disappears and toxic results soon set in if the surrounding atmosphere is dis- 
placed by hydrogen. Extreme temperatures also reverse the effect of the 
poisons. The author believes that the increased C0 2 output in the living 
tissues is a result of their battling against the poison, and that the ability to 
do battle is lost with death or with unfavorable life conditions. He concludes 
that the action of poisons on the respiration of dead tissues is a direct result of 
injury to the respiratory enzymes, while their action on living plants is indirect 
through the living protoplasm. 

Palladin believes that the increased respiration is due to an increased 
transformation of the zymogens to active enzymes, which is also accompanied 
by an increased destruction of the active enzymes. For this reason the stimu- 
lated plant shows no increase in active enzymes over the control. One teeis 
that this conclusion is not sufficiently backed by evidence. 

Palladin 6 has also attempted to ascertain the effect of contained lipoids 
upon the respiration of plant tissues. This he did by extracting dried wheat 
seedlings for two hours with various lipoid solvents (toluol acetone, benzen, 
chloroform, benzin, alcohol, etc.). In general he finds that the more lipoid 
and therefore phosphorus-containing material the solvent extracts, the more 
it reduces respiration. He believes this is related to the important role 
phosphates in zymase action. He finds, however, that the amount of reduc- 
tion in respiration is not exactly proportional to the amount of lipoid extrac e . 
He explains this in part by the fact that different solvents will extract not on y 
different amounts, but different sorts of lipoids. In substance he also says. 
"When we separate the lipoids we destroy the normal protoplasmic structure. 
We separate from the protoplasm the cement which binds its heterogeneous 
parts into a whole. I have shown that even the mechanical breaking up ^ 
the plasma structure reduces very markedly the respiration of the plant tissue. 
He points out various other means through which these solvents may a ec 
the respiration, such as the coagulation by alcohol. With other authors, 
believes the lipoids are to be considered intermediary bodies in respiratio 
that is, oxygen-carriers. He would list them with his respiratory chrom 
gens. One gets the impression that Palladin feels himself much more secu 
than he should when treading on such uncertain ground. 

6 Palladin, W., Zur Physiologie der Lipoide. Ber. Deutsch. Bot. Gese . 
28:120-125. 1910. 


Galitzky and Wassiljeff 7 have studied the effect of boiled extract of 
wheat and bean seeds upon the respiration of living and killed (by treatment 
with acetone) wheat seedlings. In agreement with various other workers, 
they find that the extracts greatly increase the C0 2 output in both living and 
killed seedlings: 60 per cent in normally acid cultures, 117 per cent in neutral 
cultures, and 86 per cent in slightly alkaline cultures. The authors raise the 
question, "Is the stimulative action due to food materials supplied by the 
extract, or to bodies of the nature of co-enzymes ? " L. and N. Iwanoff have 
assumed that the stimulation is due to the action of organic or inorganic 
phosphates on anaerobic respiration, analogous to the action of these bodies 
as co-enzymes for zymase. The authors find that peptone, glycerin, mannit, 
sodium lactate, quinic acid, sodium chlorid, and ferric chlorid have very little 
or no effect upon the respiration of the seedlings. Dextrose, saccharose, 
maltose, and sodium carbonate give a slightly increased C0 2 output, about 20 
per cent. Arabinose and ferrous sulfate give a somewhat greater stimula- 
tion of the extracts. The authors consider it especially interesting that prob- 
able intermediate products of alcoholic fermentation, lactic acid and sodium 
lactate, have no stimulative effect. This is <|uite in contrast with conceptions 
of Kostytschew. The authors are to extend the tests to various other sub- 
stances, especially to phosphates, to see whether they wall give the amount 
of stimulation shown by the extracts. One naturally wonders to what extent 
bacterial action may increase the C0 2 output. So far as methods are described, 
one certainly cannot be assured that such has not happened. — William 

Heliotropism and geotropism. — Guttenberg 8 has already shown, contrary 
to the earlier conception of most writers, that the effect of the geotropic stimulus 
is not annulled by the light stimulus, but that light of such intensity can be 
chosen that, when it strikes a horizontally placed orthotopic epicotyl from 
below, its joint action with gravity will, after various nutations, lead to the 
epicotyl permanently growing in the horizontal position. Compensation 
(placement at 45 above the horizontal) likewise results when the light of 
proper intensity strikes the vertical epicotyl horizontally. The intensity of 
light demanded to compensate gravity varies greatly with different epicotyls; 
while occasionally greater than one candle power, it is generally only a fraction 
of a candle power. Richter 9 called the results of Guttenberg into question, 


Ber. Deutsch. Bot. Gesell. 28:182-187. 1910. 


Guttexberg, H. R. v. Ueber da 


Geotropismus in parallelotropen Pflanzenteilen. Jahrb. Wiss. Bot. 45:193-231. 

J 907. 


* j ~ - — 

m us. Jahrb. Wiss. Bot. 46:481-502. 1909. 


maintaining that the results were modified by the gaseous impurities of the 
laboratory in which they were obtained. Guttenberg, 10 on repeating his 
experiments in pure air, finds essentially the same compensatory values of 
light as he found in his earlier work. Guttenberg used the seedlings of 
Avena and Brassica, forms much less sensitive to impurities than are legumes, 
with which Richter worked. Guttenberg finds in Vicia saliva, contrary to 
Richter, that laboratory air does not increase the heliotropic sensitiveness, 
but in agreement with Richter he finds the geotropic irritability lessened. 
On this point, Guttenberg's experiments are much more critical than Rich- 

ter's. — William Crocker. 

Morphology of Phylloglossum. — A recent paper by Wernham 11 represent, 
a type, at the moment becoming much too common, in which a small basis of 


neither clearly nor logically drawn. The author has examined by means of 
serial sections the anatomy of two specimens of Phylloglossum Drummondiu 
He concludes that the basal leaves of this species (the protophylls of certain 
authors) are microphyllous, although superficially relatively large in sizes 
because their traces leave the stele without leaving any gap, as is the case with 
the Lycopsida. Concerning the relation of the sporophyll traces to the vascu- 
lar system of the axis, the account is very obscure, since it is not made clear 
whether gaps are or are not present. The most remarkable feature of the 
article is the interpretation of the larger strand which passes off from the 
crown of the functional tuber toward the tuber of the succeeding year as a 

leaf trace. 


reason, as a branch supply, and the present author adduces apparently no valid 
evidence why this view of its nature should not continue to be held. On the 
basis of this imaginative interpretation, he comes to the conclusion that 
Phylloglossum was originally a megaphyllous form, which has become much 
reduced. It would be possible to prove almost anything with such reasoning 
as this. It seems highly desirable that morphologists should avoid eccentric 
conclusions of the nature illustrated by the article here reviewed. Obviously, 
conclusions of permanent value in regard to leaves or other organs can 
reached only in the case where there is no room for doubt as to the morpho- 
logical category of the structure under discussion. — E. C. Jeffrey. 

Classification of conifers.— A new classification of conifers, based upon 

morphology, geographical distribution, and geological history, is 



Guttenberg, H. R. v., Ueber das Zusammenwirken von Geotropismus un 

Jahrb. Wiss. Bot. 37:467-492. 1910. 


« Wernham, EL P.. The morphology of Phylloglossum Drummondii. Annas o 

Botany 24:335~347- figs. 8. 19 to. 



by Vierhapper. 12 It is assumed at. the outset that a group whose members 
have in common such striking characters must be monophyletic. The mor- 
phology is confined to the grosser taxonomic characters (but includes the struc- 
ture of the wood), excluding entirely the gametophytic structures, because the 
new system is based upon facts, and the gametophytes are not yet known in all 
the genera, and presumably are not yet worthy of recognition among the 
established facts. The obvious characters are analyzed and classified as 
primitive and secondary. 

The Cordaitinae are the primitive stock, which during the Carboniferous 
gave rise to the Coniferae, the Taxocupressaceae arising as an offshoot from 
Ginkgoinae, and the Abietaceae coming directly from the Cordaitinae. The 
Taxocupressaceae include the Taxoideae and Taxodioideae arising independ- 
ently from the Ginkgo stock during the Carboniferous, and the Cupressoideae 
arising during the Trias from the Taxodioideae. The Abietaceae include the 
Araucarioideae coming directly from the Cordaites stock, and the Cunning- 
hamioideae and Abietoideae arising from the araucarian stock during the 

Without commenting upon the scheme itself, it would seem to the reviewer 
that so much is now known about the gametophytes, and that so much of 
this comparatively recent knowledge is extremely significant, that it must be 
considered in any classification which claims to represent the phylogeny of 
a group.— Charles J. Chamberlain. 

Cretaceous pine leaves. — In the communication cited the authors describe 
the anatomy of a species of Finns and a supposed species of Prepinns, from the 
Upper Cretaceous of Hokkaido, Japan. z * The pine leaf, denoted by the specific 
name P. yezoensis, from the description given is not very different in struc- 
ture from the living P. Bungeana of China, since it is a soft pine with a single 
foliar bundle and apparently a three-leaved fascicle. In type it clearly differs 
from the pine leaves of the Lower Cretaceous described by the reviewer, in 
possessing a degenerate transfusion sheath, a well-marked endodermis, and 
infolded mesophyll cells. In other words, it is practically indistinguishable 
|n its general structure from the leaf of a living pine. Interestingly enough, 
in a communication from the reviewer's laboratory, shortly to appear, it will 
be shown that the wood of a pine from the American Upper Cretaceous like- 
wise resembles more nearly the secondary xylem of living pines than that 
of pines from the Lower Cretaceous. It will thus apparently be possible to 
distinguish between these two horizons by means of the nature of the pine 
flora. The Prepinus described, P.japonicus (why not P. japonica?), appar- 

ra Vierhapper, F., Entwurf eines neuen Systemes der Coniferen. Abhandl. 
K. K. Zool.-Bot. Gesell. Wien 5:1-56. 1910. 

13 Stopes, Marie C, and Kershaw, E. M., The anatomy of cretaceous pine 
leaves. Annals of Botany 24:395-402. pis. 27, 28. 1910. 


ently does not belong to that genus at all, since it is without the very striking 
internal transfusion sheath of the lower cretaceous genus, has a double 
instead of a single foliar bundle (a point of great importance, as will be recog- 
nized by those versed in the anatomy of Pinus in its living species), and no 
centripetal xylem. The authors make light of the absence of the last feature, 
but in this they are apparently not well advised. — E. C. Jeffrey. 


Conduction of stimulus. — Rothert has shown that the conduction of 
stimulus of unilateral illumination from the tip of the Avena seedling to the 
darkened basal portion occurs when the vascular strands are cut; and that a 
horizontal incision on one side, whether toward, away, or on the flank in ref- 
erence to the one-sided illumination, still permits conduction. Fitting showed 
that when the incision was away from the light and a mica plate inserted, no 
conduction occurred. The insertion of a slice of rattan in the same position did 
not prevent conduction; the latter of course allows the continuity of water and 
solution. When the mica plate was inserted in an incision on the lighted side, 
conduction was not hindered. Jensen 14 finds that with the incision away from 
the light, no conduction occurs in dry air or in water. He assumes that 
under favorable conditions the stimulus can be conducted across the wound 
while under unfavorable conditions it cannot. In saturated air the stimulus 
was also conducted from the tip to the darkened base, even after the tip (i cm. 
long) had been entirely cut off and set back and fastened by gelatin and cocoa 
butter. — William Crocker. 

Embryo sac of Pandanus. — In 1909 Campbell published an account of 
the embryo sac of Pandanus, which was reviewed in this journal (47 : 485- 
1909). In this Javanese material the fertilization stage was not secured, so 
that it was not certain that the interesting situation described is the one at 
fertilization. Now there has come to hand additional material (P. coronal us) 
which has supplied the missing stage. 15 An ordinary egg apparatus is organ- 
ized, but there occurs "a large discoidal mass of cells" at the antipodal end 
of the sac, and fusions of "polar" nuclei (up to six) were observed. The num- 
ber of cells in the sac at the fertilization stage would thus seem to be greater 
than that recorded for any other angiosperm. The amount of antipodal tissue 
suggests the situation in Sparganium, the difference being that in the latter 
genus this tissue develops after fertilization.— J. M. C. 

*4 Jensen, P. Boysen, Ueber die Leitung des phototropischen Reizes in Avena- 

kumpflanzen. Ber. Bot. Gesell. 28:118-120. 1910. 


Bull. Torr 

Bot. Club 24:293-295. figs. 6. 1910. 






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15he Armenian 


Beginning with the "Dark Ages" of Armen an 
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33 EAST 
17th ST., 









William Crocker, Lee I. Knight, and Edith Roberts 

(with six figures) 


The peg of seedling Cucurbitaceae has frequently been used 
as a marked case of adaptation to a peculiar function. It holds 
the seed coat while the elongating arms of the arch of the hypocotyl 

FlG - *• — Big Tom, showing the pegs functioning in the removal of the coats 

withdraw the cotyledons from the coat 
the peg functions is well shown in fig. I. 

The method by which 



Mirbel 1 and Tittman 2 very early showed the presence of this 
organ throughout the Cucurbitaceae. While it is well developed 
in the epigean f orms, it is very rudimentary in the hypogean forms, 
such as Megarhiza calif ornica and Sicyosperma gracilis. The 
organ is by no means limited to the Cucurbitaceae, but appears in 
various genera of a number of families: Mirabilis, Oxybaphus, 
and Abronia (Nyctaginaceae), Martynia (Martyniaceae), Lind- 
heimera (Compositae), Mimosa (Leguminosae) ,. Tribulus (Zygo- 
phyllaceae), Eucalyptus (Myrtaceae), Cuphea (Lythraceae), etc. 
While in Cucurbitaceae the peg appears only after germination 
has progressed considerably, in other forms, as Eucalyptus and 
Cuphea, it is already laid down as a complete ring in the mature 
seed, and with germination completes its development. 

Tittman recognized the biological significance of this organ in 
the Cucurbitaceae, and showed that it appears only on the lower 
side of the developing hypocotyl. 

Tscherning 3 gives a rather full description of its histology 
and physiology. He states that it is a parenchymatous out- 
growth. While the greater diameter of the parenchymatous cells 
in other parts of the hypocotyl is longitudinal, in the peg zone it 
is radial. The number of layers of cortical cells is also somewhat 
greater at the peg zone. Tscherning describes the effect of the 
position of the seed during germination upon the development and 
functioning of the peg. He says that when the radicle points 
vertically downward the swelling does not occur, and the coty- 
ledons push above the ground still bearing the coat. With radicle 
pointing vertically upward, the peg develops on the concave side 
of the arch, but does not remove the coat. When the seeds are 
planted on edge, the peg develops on the concave side of the arch 
and wedges between the two valves of the coat, thus succeeding in 
removing a considerable percentage of them. When the seeds are 
planted on a flat face, the peg develops on the concave side of the 
arch and attaches itself securely to the lower valve of the coat, 
thereby insuring its removal. Tscherning emphasizes the deve - 
opment of the peg on the concave side of the arch, and speaks o 
it as a lateral pushing out of the cells due to the inhibition in the 

*» ». J See under Noll, footnote 8. 


elongation on the concave side of the hypocotyl. While his 
relating its place of development entirely to the arch is probably 
correct, as we shall see later, his view as to how the arch brings 
about such a placement is crude. We shall see also that his 
claim that the peg does not develop when the seed grows with the 
radicle pointing vertically downward is incorrect. 

Flahault 4 claimed that seedlings that developed when the 
removal of the coats occurred in the normal way were far superior 
to those in which it was prevented by breaking away a portion of 
the lower valve of the coat so that the foot could not get a hold. 
This advantage he attributes of course to the hindering of assimila- 
tion by the retained coats. He claimed that the peg develops 

at any point on the hypocotyl necessary to enable it to hold the 

Charles Darwin, 5 contrary to Flahault, found that the peg 
develops only at the zone between root and stem. The lower 
face is root, as shown by the presence of root hairs and reaction 
to potassium permanganate, while the upper face is stem. Darwin 
described a number of experiments, similar to those of Tscher- 
ning, testing the effect of the position of the seed upon the develop- 
ment and functioning of the peg. While Darwin clearly showed 
that the peg is located at the border between root and stem, he did 


not show the stimuli involved in its lateral placement. 

Francis Darwin 6 later showed that seedlings of Cucurbita 


rotating clinostat with 

a horizontal axis, gave pegs completely surrounding the hypocotyl 
and approximately equal on all faces. He 


that gravity 



effective on the clinostat and only equalized in its action on the 
several flanks of the exposed object. The conclusion concerning 

4 Flahaij] 
Soc. France 24 

5 Darwin 


102-107. New York. 1892. 


Cambridge. 1895. 



the nature of the effect of the clinostat is correct, as shown by 
Fitting 7 and others, but the assumption that this experiment 
proves it is quite another question, and, as we shall show later, 
quite out of accord with a number of other facts. 

Noll 8 mentioned the fact that Francis Darwin's conclusion 

stimulus in determining 

meager experiments. He 


on a clinostat produce ringlike pegs, while the remainder show 
sharp arching, with a one-sided peg. Noll stated that all flanks 
of the hypocotyl are qualitatively equally capable of peg-develop- 

peg. He 

stimulus to peg-development, for the peg still appears when the 
coats are removed. The contact of the coat, however, increases 
the size of the peg. He performed a number of experiments in 
which seeds were germinated with the two opposite broad faces 
exposed to gravity alternately and for equal periods. In all these 


^~^ ^^^^ ^^"^ ^W ^'— w ^fc- ■ ™ -^fc- ^^^ ^^r W « ^^^^ ^^^ ^H^H ^^_* ^k ^ k # % I ^fc J Vf ^L f j| ^ # 1 W ft. \ ft ■* ^ h T ^ ft 1 ft ■ W m ft m A ft ft J ^fc__f ^^^^ ^B^k ^ ■ ^ _^h 

emphasized these as showing that the peg stands in strong rela- 
tion to gravity and owes its stimulus for formation to gravity. 
Many experiments were also performed in which seedlings were 
grown with the radicle directed perpendicularly downward or at 

from this Dosition. From 

that the gravity stimulus reaches beyond the lower pole of the 
seedling axis. At a deviation of 5 from the vertical he found a 
weak peg on the upper side, while at 7^5 the peg was almost 
exclusively on the lower side. He fixed the limit of the field of 
stimulation at 6°-8° deviation from the vertical. He specified 
that these are to be taken only as approximate figures and as 
applying to Cucurbita Pepo. In two seedlings slight pegs were 
found on the upper side at a deviation of 8°. Seedlings were also 




— „ ^v,„w^ UV peg, UUt 11C U1U 11WU »*«,« 

On account of great variation in response, Noll empn - 

» Fitting, Hans, Jahrb. Wiss. Bot. 45:575-600. 1909. 

•Noll, F., Zur Keimungs-Physiologie der Cucurbitaceen. Landwirt. Ja r • 
30:145-165. 1901. 


sized that reliable conclusions can be drawn only by the employ- 
ment of a large number of seedlings in each culture. 

It is evident that both Francis Darwin and Noll considered 

gravity as well as the arching of the hypocotyl as direct stimuli 


Noll says (p 

inseitiee Ausbildune des Wulstes 

nis zweier heterogener Reize ein. Die localisierte Entstehung 
des Wulstes ist einerseits abhangig vom Gravitationsreiz. Der 

Wulst bildet sich auf der jeweiligen Unterseite Die ein- 

\ Wulstbildung wird andererseits auch bedingt durch die 



mainstay for the memory theory of plant response. In this, of 
course, he assumed that its development and position are directly 
determined by gravity. Whether gravity acts as a direct stimulus 
to its production and placement is the principal question to be 
tested by the following experiments. 

Methods and materials 

Noll failed to control the factors involved in this problem in 

him to determine 

in the lateral placement of the peg. He apparently failed to notice 
the potency of contact of the coats on the one hand, and of the media 
on the other, in arch-production. In the experiments here given, 



different inclinations, were freed from 


cotyledons were clasped at the central region. The whole appa- 
ratus was kept in the dark and watered by a very fine spray heated 
to 23 C. The spray was formed by forcing the water by means 
of tap pressure through a tank of considerable size kept in a water 
bath at constant temperature, and allowing it to break against a 
plate of glass. In this way contact of both the coats and the soil 
media is entirely eliminated. Contact, as data later given will 

'Darwin, Francis, New Phytologist 5:199-207. 1906. 




show, is a very important influence. We are unable to se 
Noll could get results at all dependable without taking thi 
caution. He was always studying the effect of two stimuli 
his results purport to be considering one. 

The seeds used in this work were obtained from Vauc 
will be mentioned by the trade name used by that dealer. 

Experiments and discussion 


As has been stated, the contact of coats plays a very important 
part in arch-production, and therefore indirectly upon the lateral 

Fig. a— Crop of Hubbard squash grown in spray at 23 C; coats removed at 
the tip and radicles pointing downward. 

placement of the peg. This effect was tested in two ways : (1) by 
growing seeds upon a clinostat with coats intact and with coats 
removed at tips, and (2) by a similar growth of seeds held between 
cork strips with radicles pointing downward or approximately so. 
These were kept in a dark chamber and watered with a spray 
at 23 C, as described above. There is some variation in differ- 
ent varieties in the response to contact. For example, the pump- 




kin called Big Tom, especially when grown on the clinostat, gives 
a somewhat higher percentage of strong arching in response to coat 
contact than does the Hubbard squash. Fig. 2 shows all the 
individuals of a culture of Hubbard squash grown as described 
above, coats removed at tip, and radicles pointing downward. 
Of the 26 seedlings, none are sharply arched, and all have pegs 

either two-sided or ringlike. 


pegs on one side than on the other. It is evident then that in 

Fig. 3.— Crop of Hubbard squash as in fig. 2, but coats not removed at tip; shows 
sharp arching induced by contact of coats. 

the HuhharH eniiacii little ai-r4Wnfr qnrl nn nnp-sided De£s are pro- 

downward, and with no contact of either coat or media. Fig. 3 
shows a similar culture except that the coats were intact during the 
growth. Of the 42 seedlings, 29 are sharply arched and 13 very 
slightly or not at all. This shows contact of coats to be very potent 
in arch-production. Of those that are sharply arched, 8 have 

*■ « m. m 1 



In this case the 


ching is against gravity, which em 


tact of coats as a stimulus to arching. By a comparison of figs. 2 
and 3, one can see clearly the effect of contact upon the size of the 


A culture of pumpkin (Big Tom), with coats removed, radicles 

pointing vertically downward, and grown in a spray as described 

above, produced 39 seedlings. Of these none were sharply arched; 

19 showed very small two-sided pegs; 19 were essentially pegless; 
and 1 showed a small one-sided peg on the concave side of the 
slight arch. A similar culture of the pumpkin (Big Tom) with 
coats intact gave 31 seedlings; of these, 25 were sharply arched, 
with pegs entirely on the concave side of the arch; 1 was little 
arched, with peg on concave side of arch; 3 were pegless; and 2 
had equal, small, two-sided pegs. From these data it is evident 
that in the pumpkin (Big Tom) contact of coats is even more 
effective in producing arching than in the squash (Hubbard), 
and unlike the case of the squash, the arching shifts the peg entirely 
to the concave side of the arch. From the large number of peg- 
less seedlings appearing in the culture with the coats removed, it 
is also evident that the arch not only determines the lateral place- 
ment of the peg, but in some cases even its appearance. One may 
be inclined to think the existence of the peg is perhaps determined 
by the contact of the coat, rather than by the arching produced 
by the coat; but it is not probable, for of the 6 not arched in the 
culture with coats on, 3 are pegless. It is apparent, therefore, 
that a large number of the pumpkins (Big Tom) are pegless if 
developed without considerable arching, and that arching deter- 
mines in about half the cases whether or not there will be a peg 
as well as the placement of the peg on the concave side of the arch. 

Fig. 4 shows a culture of pumpkin (Big Tom) with coats removed 
at the tips, and grown on the horizontal clinostat in a spray. Of 
the 30 seedlings in the culture, none are sharply arched; 14 have 
small two-sided pegs ; 8 show slight pegs on the concave side of 
the slight arch; and 8 are essentially pegless. Fig. 5 shoWS a 
similar culture except the coats are intact. Of the 24 seedlings, 

20 are sharply arched, with pegs entirely on the concave side of 
the arch; 2 are slightly arched, one of which has an equal two- 
sided peg and the other a slight one on the concave side of the arch; 




3 2 9 

Fig. 4. — Crop of Big Tom grown on a clinostat in a spray; temp. 23 ; coats 
removed at the tip. 


Fig. 5. — Crop of Big Tom grown as fig. 4, except coats were not removed at the 


2 show no arch and are essentially pegless. One is struck here 
by the small percentage of pegless seedlings in the culture in which 
the coats are removed, as compared with corresponding seedlings 
grown with the radicle pointing vertically downward. This is 
easily explained, however; the arching does not have to take 
place against gravity, due to the equalization of this stimulus by 
the clinostat. In short, the autonomic disposition to arch is unre- 
strained. This fact, with the fact that slight arching produces 
a one-sided peg in this form, makes the data exactly what must 
be expected. Again, one sees that the contact of the coats is 
much more effective in producing arching and one-sided peg- 
development than in the culture in which the seedlings developed 
with the radicle pointing vertically downward. This again must 
be expected, for the arching does not have to occur in opposition 
to gravity, since this stimulus is equalized by the clinostat. 

Noll states that more- than 50 per cent of pumpkins grown on 
a horizontal clinostat show sharp arching, with the peg entirely 
on the concave side of the arch. This is the case if one allows two 
factors to act at once : contact of coats and the autonomic tendency 
to arch. But Noll's conclusion that such results will follow from 

tendency to arch alone is erroneous, and was pos- 


important influence 

contact of coats in the arching. Darwin's statement 


, the peg is 

almost equal on all sides, certainly holds for the squash (Hubbard), 


in our 



evident. Contact of coats is an extremely im 

the pumpkin and the squash. It 
produces strong arching even against gravity. In the pumpkin 
(Big Tom) any considerable arching increases greatly the percent- 
age of seedlings that show pegs ; and it causes the development of 
the peg on the concave side of the arch. In the squash (Hubbard) 

arching is far less effective in determining the latere 
the peg. The autonomic tendency to arch is very 

,pkin (Big 


Tom), as shown by the clinostat experiments where the coats 
are removed. It is also evident that results obtained without 
reference to the effect of contact of the coats cannot lead to a 
knowledge of the part played by the several factors in determining 
the lateral placement of the peg. 


As has been stated, Noll claimed that seedlings grown in a 
vertical position, with the radicle pointing downward, gave two- 
sided or ringlike pegs. He found, however, that a slight deviation 
from this position (8? 5) produced only one-sided pegs. We ran 
numerous experiments on various varieties of cucurbits for the 
purpose of testing this statement. The results are tabulated in 
the accompanying table. "Vertical" in this table means that the 
seed was held so that the plane passing through the largest dimen- 
sion of the seed was in a vertical position and the radicle was point- 
ing downward. Deviations from this plane were made by tilting 
the plane of the seed the number of degrees from this position that 
is recorded in the table. The tilting always exposes one face 
rather than the edge of the seed to gravity. The seeds were held 

in position by two pieces of cork lightly clasping them at the central 
region. The whole culture was grown in darkness in a spray, so 
that no contact of media was involved and no contact of coats 
except when specified. 

A glance at this table makes it very evident that it furnishes 
no ground for Noll's conclusion that a deviation of 
gives only one-sided pegs. Our extensive experiments 
unable to understand how he obtained such results. 

over 8? 5 



placement of the peg when the coats are intact than when they are 
removed at the tip. This is due to the fact that the arching is 
made much sharper by the coat contact, and the lateral placement 

• f 



varieties in the effectiveness of the deviation for the lateral place- 
ment of the peg. In Big Tom it is most effective, while in Boston 




























































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marrow it is least so. In the former no two-sided pegs appear 
at 13 5 deviation or above, while the latter at 180 deviation still 
shows about 8 per cent with two-sided pegs. 

In the exposures with the coats removed, one can see the effect- 
iveness of gravity alone in the one-sided peg-production. It 
must not be forgotten that the effect of gravity in lateral placement 
of the peg is probably entirely indirect and acts through the pro- 
duction of an arch. 

As has been mentioned, Noll thought that not only the develop- 
ment of the peg but its lateral placement is called forth by the joint 
action of two heterogeneous stimuli: gravity, which causes the 
development on the lower side of the hypocotyl; and " organ- 
form" stimulus, which causes the development on the concave side 
of the arch. Noll compared the second of these to the appearance 
of lateral roots on the convex side of a curved main root. His 
main evidence for the conclusion concerning two heterogeneous 

-** I*** « - * * . - . - - ••»• M 

his experiments in which he grew seedlings 


a one-sided peg were true, his conclusion would be entirely justified. 
Our results, derived by the use of five different varieties of cucur- 
bits, of very different characters so far as peg-development is con- 
cerned, and obtained with greatest care in eliminating all factors, 
except gravity, show that it takes a deviation of not merely 8° 5, 


marrow of more than 135 , to insure a one-sided peg. 




the lower side only, and yet our results show that many seedlings 
thus grown produce two-sided pegs. Noll's two heterogeneous 
stimuli are not adequate to explain these results. This 


becomes yet less in 
Noll assumed 


' in inducing peg-development on the upper side of the hypocol 

lie deviation from the vertical is over 8? 5. 

It has alreadv been pointed out that in Eucalyptus and Cuph 




a ringlike peg is laid down in the formation of the seed, and with 
germination this enlarges somewhat. In these 


development a 


alone by external 

stimuli. We find that such a ringlike De£. rather small 




Vic 6. — Pegless seed- 
lings of Big Tom. 

^arkinsonia). In this form the arching of the 
marked and the peg does not function in the 
)at; likewise, external conditions do not seem 
1 placement. Results already stated, as well as 

data to be stated later, lead us to believe 
that the peg of the cucurbits is, to a con- 
siderable degree, a natural outgrowth of the 
seedling, and that it is approximately equal 
on all flanks if arching is avoided. We must 
of course accept the quantitative greater 
development on the two broader faces of 
the hypocotyl over the two narrower. It 
is also evident that in different species the 
size of the outgrowth varies greatly; it is 
relatively large in squashes (Hubbard, Bos- 
ton marrow, and Calhoun),. and relatively 
small or even absent in pumpkins (Big Tom 
and Sugar pie). We have also found the 
latter to be the case in various watermel- 

ons and cucumbers. 



avoided, different species will show all the 


have very prominent ringlike pegs. We 
the contact of coats, as figs. 2 and 3 show, greatly increases the size 
of this outgrowth. The size is also greatly increased, at least on 
one flank, by arching. 

- w — — 

stimuli cause the development of this 
assumed, but rather what stimuli 1 
and tend to increase its size. 



It is certainly evident, as all who have worked on this subject 

agree, that arching or " organ-form " stimulus, as Noll termed it, 

leads to the shifting of the peg to the concave side of the hypocotyl, 

or, as it would be better termed, to the development of the peg 

on the concave flank of the hypocotyl. Both gravity and contact 

of coats aid in arch-production. Of these, as our results show, 

the latter is much more important. Contact of coats also greatly 

increases the size of the peg. Other questions that should be 

answered in this connection are whether or not gravity acts as a 

direct stimulus in the lateral placement of the peg, and whether 

or not it tends to increase the size of the peg on the flank exposed 
to it. 

As has been repeatedly said, both Darwin and Noll assumed 
that gravity is a direct stimulus, not only in determining the 
existence of a peg but also its lateral placement. Darwin's evi- 
dence for this was that a ringlike peg is found when the seedlings 
of Cucurbita ovifera were grown in a horizontal position on a slowly 
rotating clinostat. Noll's evidence for this conclusion included 
two other facts: (1) seedlings with the radicle pointing downward 
but deviated more than 8? 5 from the vertical gave pegs only on 
the lower side, and (2) seedlings turned over every few hours 
during development gave pegs ringlike or two-sided. We have 
shown that Noll's deviation experiments do not at all accord with 


one factor at a time. As for the clinostat and turning evidence, 
it is just what would be expected if the peg is an outgrowth approxi- 
mately equal on all sides, if sharp arching is avoided. Of course 

from time to time 

ing, and thereby give the natural ringlike or two-sided development. 

Further evidence as to whether gravity is a direct stimulus in 

determining the existence of a peg, its lateral placement, or its 



A large number of cultures of the several varieties of cucurbits 
worked with were made on a centrifuge with a vertical axis. The 
acceleration varied from two gravities to eighteen gravities. The 


cultures were grown in a spray at 23 C; the coats were removed 
to avoid all contact; and the long axis of each seed was arranged 
parallel with a radius of the centrifuge, the radicle directed away 
from the axis. 

Three facts were noticeable in such seedlings, especially in those 
grown with the greater centrifugal acceleration: (1) the hypocotyls 
were very straight, (2) the pegs were smaller than in similar cul- 
tures on clinostat or with radicles pointing vertically downward, 
and (3) the pegs though small were approximately equal on all 

If, as Noll assumes, gravity calls forth the peg in seedlings 
with the radicles pointing downward, one would expect larger 
pegs with increased gravities, unless the rather remarkable situa- 
tion exists that one gravity is the optimum or is greater than the 
optimum for peg-development. But how can one account for the 

and very regular peg? This is undoubtedly due 

more meager 


by the centrifuge, which 

t must not be forgotten that a centrifug 

mass accelerations at riant ancrlfQ tn eai 

one due to 


in the direction of the pointing root, and one gravity downward. 
If a flat side of the seedling faces downward, and the machine 
gives a centrifugal acceleration of three gravities, three gravities 
act in the direction of the pointing root and one gravity downward. 


j o - — — "■ j — ' , • 

peg-development on the lower than on the upper flank. If this 
organ were sensitive to gravity, one would expect it to show a 
greater response on the side where one extra gravity was acting, 
especially when the centrifugal acceleration was only three gravities. 

Of rnnrsA it io anti^U, — „„:ui~ x.i__x nr^„^^v 1„«r onnlies in SUCh 



r . T „ 1( . luUi! «.uai luc application 01 one gravity 0.3 a ^«.*~ 

not be perceived when three gravities are acting at right angles 
to it. In geotropism of Vicia Faba epicotyls, however, Weber S 
law applies in the proportion of 24-25, that is 4 per cent exceSS 



It must be remembered that while this experiment does not prove 
that gravity is not a direct stimulus in determining the size and 
lateral placement of the peg, it certainly furnishes no evidence 
that it is. 

A detailed statement of the results of one experiment with Big 
Tom on a centrifuge with a horizontal axis will suffice to indicate 
the nature of the results obtained with that apparatus. Here, 
also, the coats were removed at the tip and the temperature kept 
at 23 C. by means of a spray. The revolutions were 440 per 
minute and the circles 3 and 6 inches in diameter. The 3-inch 
circle gives 8.25 gravities, while the 6-inch gives 16.5. In the 
3-inch circle 15 were essentially pegless and 2 with slight two- 
sided pegs. In the 6-inch circle 25 were essentially pegless, 4 
with slight two-sided pegs, and one with a peg toward one edge 
of the seed. The marked tendency to show pegless forms is 
manifested here, as in all experiments on the centrifuge, with three 
or more gravities. It seems that these accelerations prevent all 
disposition to arching and to arching nutations; hence they give 
many pegless forms. 


A number of cultures were made on the oblique clinostat as 
devised by Fitting. 10 The cultures were watered by a spray at 
23 C. and grown with coats removed at the tip to avoid contact. 
The end of the clinostat axis to which the frame for bearing the 
seeds was attached was pointed upward from the horizontal, while 
the radicles of the seeds pointed toward the axis. This insures, as an 
understanding of the oblique clinostat will show, that extreme arch- 
ing is avoided. The axis was placed at various angles with the hori- 
zontal, and the large plane of the seeds at corresponding angles 
with the axis. When the axis varied 5 from the horizontal, and 
the large plane of the seed was broken 5 from the axis, at one 
Point in each revolution one flat face of the seed was exposed to 
gravity at an angle of 8o° from the vertical (radicle pointing down- 
ward) and the other face at oo°. As soon as considerable growth 
occurs, of course geotropic response occurs in favor of the 90 
exposure, and this throws the flank of 8o° exposure more nearly 

'• See footnote 7. 



9q°, thereby avoiding sharp arching. The difference of exposure 
is marked then only in the early stages, and other exposures with 
qo° against 76 and 90 against 85 gave no peg-development in 
favor of the 90 either in the pumpkin or squash. It has been 
shown that the effectiveness of geotropic stimulus in orthotropic 
organs is approximately proportional to the sine of the angle, and 
therefore the 90 exposure is stronger than any of the others. If 
gravity is effective as a form stimulus in causing the lateral place- 
ment of the peg, we might expect it to be manifested by a larger 
peg-development on the flank exposed at 90 . These experiments 
give no indication of it, and yet it would hardly be expected that 
such differences in exposure would show an effect, since much 
greater differences failed to do so in the centrifuge experiments 
described above. It must also be pointed out that, unlike the 
centrifuge experiments, these experiments are rather unsatisfac- 
tory because the arching occurs in favor of the 90 , and that factor 
is sufficient, if great, to produce lateral placement of the peg. On 
this account, only angles differing rather slightly can be compared. 
It is evident, however, that these experiments, as do the centri- 
fuge experiments, furnish no evidence that gravity is a direct 
stimulus to the lateral placement of the peg. 

and summary 

As one sees from the experiments given above, there is no 

evidence that gravity acts as a 


»/ the peg. Certainly, then, Darwin is not justified in using this 
assumption as a main prop to a theory (mnemic theory) which 
itself looks away from rather than toward progress in the knowledge 
of plant response. 

Assuming that all arching is avoided, the following facts seem 
to hold: the peg is to a considerable degree a natural integral 
part of the plant; it develops on all flanks of the hypocotyl approxi- 
mately equal (granting perhaps that it is somewhat larger on the 
broad flanks in many of the cucurbits) ; it varies in size from the 
very slightest outgrowth appearing in a small percentage of Big 
Tom to the large pegs of the Hubbard squash. 

It may be laid down with the formation of the seed, as m 


Eucalyptus and Cuphea, where it is an equal ring all around the 
organ; or its formation may begin after germination, when its 
position and size are determined by the factors we have shown to 
be effective. 

The lateral placement is apparently brought about by the 
arching of the hypocotyl. Two stimuli aid in the formation of 
the arch: contact of the coats and gravity. Contact of the coats 
is by far the more effective, for it will induce very sharp arching 
even against gravity. 

In forms like the Boston marrow, gravity independent of contact 
with a deviation of 170 from the vertical gives strong enough 
arching to produce only 90 per cent with one-sided pegs. In 
Big Tom and various other forms it is somewhat more effective. 

Arching leads to an increased development of the peg, as well 
as to its lateral placement, and in many cases it produces a peg 
where it would not otherwise appear, as in Big Tom. Contact 
likewise increases the size of the peg independent of its effect 
through arch-production. 

No evidence obtained from this detailed study indicates that 
gravity, as a direct stimulus, in the least increases the peg-develop- 

The University of Chicago 



Frederick H. Blodgett t 

(with plates viii-x and seven figures) 


The genus Erythronium includes some fifteen species of bulbous 
plants of the north temperate zone, one of which occurs from the 
Pyrenees, across France, southern Europe, and Asia, to Korea and 
Japan. The remaining species are North American. 

The plants are perennial through the annually renewed bulbs. 
These are developed at gradually increasing depths from the seed- 
ling to the flowering stage, after which they are formed at a nearly 
constant depth in the soil. The bulb consists of a few thick scales 
surrounding the stem apex; the aerial structures push upward 
from the base through the cavity inclosed by the inner scale. The 
first aerial structures are single leaves, two leaves and flowers 


appearing only after several years. 

Special details of the development of one or another of the 
species have been examined by several writers; but some stages 
have been omitted in these accounts, and in others some obscurity 
exists as to the exact nature of the structures mentioned. A con- 
secutive account of the development of the plant, with special 
reference to the origin and structure of the stem apex, and its 
outgrowths, from its inception in the embryo to the final formation 
of the flower bud, is the subject of this paper; which is thus a study 
of the vegetative development of the sporophyte, consideration of 
the gametophyte being omitted. 

Erythronium americanum is taken as the basis of the study, 
since material of this species is abundant in the vicinity of Balti- 
more, and especially since it shows more specialization in n=> 
vegetative development than do the other forms. Comparisons 
have been made with other species, as will be noted, and the 

« Contribution from the Botanical Laboratory of the Johns Hopkins Universit} 

no. 14. 

Botanical Gazette, vol. 50] 



differences found have a probable bearing upon the evolution of 
the genus. 


thickness; except in the case of the hard seeds, no special methods 

were necessary 


his thanks for helpful criticism and maintained 

the progress of the work. 


The flowers of Erythronium americanum appear during the first 
ek of April in this region, and the seeds ripen early in June, 
m which seedlings arise the following March. The main body 


chalaza and a fleshy raphe nearly double the bulk of the seed, 
subject to individual variation. The 



entiated (fig. 1). 


seed becomes moistened by the fall rains, and the embryo begins 
to elongate. The growing embryo enlarges first to occupy the 
space filled by the spongy endosperm, in which it is imbedded, 
ordinarily by the first week of October. The tip of the cotyledon 
is organized into a haustorial organ, by which the hard endosperm 
is absorbed. Elongation proceeds slowly during the fall, so that 
the embryo is half the length of the seed in December (fig. 2). 
The rate of growth seems to be closely related to the abundance 
of moisture in the soil. The tip of the radicle is pushed about the 

>pyle. The stem apex at this 
stage appears in a narrow cavity extending from the surface nearly 
to the center of the tissue just behind the radicle. The hypocotyl 
does not elongate, and is practically absent. As the embryo con- 
tinues to elongate, the stem apex is carried forward, and retains 
a constant position in relation to the radicle during the elongation 
of the cotyledon (fig. 3). The zone of elongation during 
is in the lower part of the cotyledon, just above the cavity in which 
the stem apex is situated. Growth is nearly vertical, and after 
penetrating the soil 1-3 cm. the descending axis comes to rest. 




When the endosperm has been exhausted, the elongation of the 
descending axis ceases. The zone of elongation in the cotyledon 
is now located near the upper end of the cotyledon, so that the 
tip of the cotyledon is withdrawn from the empty testa, and 

as in Allium (Sachs 18). From the 

ght, much 

haustorial cells at the tip of the cotyledon two vascular bundles 
extend downward to the base of the root, just below the level of 
the stem apex, as in Tulipa (Irmisch ii). In some species (E. 
grandiflorum, and E. Hartwegii) there are three cotyledonary 
bundles (Sargant 19). After reaching the light, the cotyledon 
attains a total length of 8-10 cm. above the soil line (text fig. 6), 
and the elbow formed in the withdrawal from the seed coats 
gradually straightens. The cotyledon is cylindrical below, but may 
be considerably flattened in the upper portions, the two bundles 
lying side by side (figs. 4, 5). 

The apical dome becomes differentiated during the descent of 
the stem apex, and the cavity in which it is inclosed changes into 
a curved pocket by the elongation of the walls of the cavity (figs. 
6-10). The apex then rises as a 

dome from 

pocket. The space about the apex has a depth about equal to 

g axis comes 

The thickness 

of the walls decreases from the axial to the opposite side, which 


lies immediately below the opening from the inte 
to the exterior. The base of the cotyledon thus 
sheath about the stem apex. The radicle has as yet grown but 
little; soon, however, the primary root is organized and advances 
from the end of the descending axis (text fig. i). The primary 
root is the only one in the life cycle which responds positively to 
gravitation. At the base of the root a considerable cushion is 
formed, upon which root hairs are produced abundantly. By the 
growth of adjacent tissues the base of the root becomes oblique, 
and seems to be fused to the side of the cotyledonary sheath for 
a short distance (figs. 6-9). The stem apex, or plumule, has so 
far no vascular supply from the rest of the seedling, the cells below 
the apical dome being meristematic and undifferentiated as yet 
into tissues. The plumular trace is inserted upon each of the two 
cotyledonary bundles, just before they unite to form the root 





in length (figs. 

16-20). This theoretically constitutes the "hypocotyl" in these 
seedlings (S arg ant 19). 










Fig. i. — Diagram showing development of seedlings: 1, descending axis during 
first stage; 2, inception of droppers and development of root; j, dropper actively 
elongating downward, second stage of descent. 

First vegetative period 

The cotyledon functions as the first foliage leaf, and provides 
material for the further descent of the stem apex. Abundant 
stomata and large intercellular spaces are present in the body of 
the cotyledon; moreover it has the power of recovery from wilting 
to a high degree. This was found by allowing the soil to become 
dry and then watering, when seedlings which had become prostrate 
recovered their erect position, unless the cells had actually been 
killed. The cells nearest the tip were the first to show the effects 
of too great drying. 

Soon after the cotyledon becomes active photosynthetically, 



and the primary root is protruded into the soil from the end of the 
descending axis, the further descent of the stem apex begins. The 
sheath in the base of the cotyledon, in which the apical dome is 
located, elongates by the proliferation of the cells in the walls of 
the sheath so that the base of the cavity is lowered in reference to 
other points. As the structure so formed is positively geotropic 
in its growth response, it may conveniently be called a " dropper. 
The apical dome, being inserted at the base of the sheath, is carried 
forward in the elongation as a terminal bud (figs. 10, 13). 

The dropper originates from the cotyledonary sheath through 
the irregular distribution of the dividing cells in the walls of the 
sheath. The base of the sheath is displaced laterally by the multi- 
plication of cells between the apical dome and the axis of the 
cotyledon (figs. 6-8). This pushes the base outward from its 
original position. While the displacement is proceeding in the 
lateral direction, an elongation is also taking place, so that the 
dropper begins to push forward into the soil during its displacement. 
Soon after the axis of the dropper becomes established as a direction 
of growth, the zone of elongation becomes confined to the region 
immediately adjacent to the base of the apical dome. This places 
the growing tissue close to the apex of the dropper, as if it were a 

losing walls of the dropper do not grow equally, the 




of the sheath cavity. The stem apex thus becomes located on the 
side wall, instead of at the base of the sheath (fig. 13). The axis 
of the dome itself may become almost horizontal through this 
displacement. The oblique position of parts inaugurated at this 

ted bulbs of 
;he stem aoe: 

The lateral loca- 

from the aDical dome 

The apical outgrowths push upward into the cavity about the stem 
apex, and thereby appear to be at nearly right angles to the axis 
of the dome (fig. 13). The rate of growth of the apical dome and 
that of the developing scale is so nearly identical that the dome 


itself is lost as a distinct group of cells. When the scale has so 
far developed that the margins of the leaf rudiment meet in front 
of the median line of the scale, then the apical dome becomes 
distinguishable again. The dome then forms a low mass of cells 
projecting from the face of the axial side of the scale into the space 
inclosed by the united margins. Continued growth soon makes 
the dome more conspicuous, and at the same time readjusts its 
position to a more central point (figs. 10, 12, 13). The tissues of 
the scale rudiment and of the stem apex are nearly alike in their 
staining qualities, and continue to be meristematic until a con- 
siderable size is reached. Thus in the first stages of scale formation, 
the growth in length of the apical dome and the growth in thickness 
of the scale rudiment are so nearly equal that there is no difference 
in staining reaction to aid in separating the two morphologically 
different tissues. It is not until the scale leaf has inclosed the 
apical dome by the forward growth of the margins of the scale, 
that the dome is to be separately recognized. This obscuring of 
the apical dome appears to be a recurring feature in the normal 
development of the bulbs, as the successive bulb scales are formed. 

The dropper grows downward to a distance varying from a 
few millimeters to 4 (rarely 5) cm.; in most cases the dropper is 
about 2 . 5 cm. in length. This growth is accomplished during the 
period of activity of the cotyledon as a leaf, by which the needed 
starch and other materials are elaborated. At the end of three or 
four weeks of development, the apical bud within the dropper 
sheath becomes enlarged by the deposit of storage starch, and 
the stem apex with its inclosing bulb scale becomes the primary 
bulb (fig. 14). The seedling dies from the apex of the cotyledon 
backward, gradually involving all tissues except the bulb, which 
thus becomes isolated in the soil. The withered walls of the drop- 
per sheath form a husk about the bulb. In the axil of the dropper 
sheath a bud is formed, which rarely develops. 

During the active descent of the dropper, vascular connection 
from the stem apex to the base of the cotyledon is maintained 
through the bundles in the axial wall of the dropper. This tissue 
is to be regarded as cauline in character, since it terminates in the 
growing point of the stem. In its earliest stages the vascular 


strand appears as a few elongated cells between the base of the 
apical dome and the base of the cotyledonary bundles (figs. 8-10, 
16-20); as the dropper elongates these cells increase in number 
by additions at the growing end, and become more completely 
differentiated into vascular elements (figs. 8, 11, 13). At the 
end of the strand branches are given off into the scales developed 
from the apex. The vascular elements are only slightly developed 
in the bulb bundles, since there is little strength required, and the 
bundles are surrounded by abundant parenchyma. The spiral 
vessels are the most easily distinguished parts of the bundles. 

During the summer the bulb shows no external signs of activity, 
but the stem apex is then organizing the first foliage leaf for the 
second vegetative season. In its inception the leaf resembles a 
scale leaf, but soon becomes differentiated into a blade and a basal 
portion. By the elongation of the base, the blade is elevated 
above the stem apex (fig. 15, I). The apical dome is thus left in a 
position simulating that of an axillary bud, with the leaf as its 
subtending organ. But the further development of the dome, 
as well as its immediately preceding history, identifies this as the 
stem apex. The leaf and the apical dome are in the relative devel- 
opment given about July 1. 

In the development of the foliage leaf, the elongation of the 
petiole lifts the lamina above the stem apex before the latter 
becomes inclosed by the development of the margins of the blade. 
The margins grow outward from the median portion of the leaf 
rudiment, their edges passing each other above the level of the 
stem apex. The base of the petiole grows about the stem apex, 
so that this becomes inclosed in a sheath much as was the apical 
dome in the cotyledonary sheath. A small opening connects the 
cavity of the sheath with the space outside the petiole in the bulb. 

At this time there appears in the axil of the leaf an axillary 
bud, which from its position in reference to the apical dome seems 
to be borne upon the surface of the dome, rather than to be truly 
axillary to the leaf. A bud is also developed in the axil of the inclos- 
ing scale. The stem apex develops usually two bulb scales during 
the late summer, thus forming a bulb rudiment at the base of 
the leaf by the close of the year. In these primary bulbs the stem 


apex alone develops into a bulb in most cases; but in older plants 
the axillary buds may also develop into bulbs. When the bulbs 
of any age are renewed in situ, the apical bud forms the single 
new bulb; but when there are runners developed, the axillary buds 
also form bulbs, being then the terminal buds of runners which 
become isolated in the soil as in the case of the primary bulb from 
the dropper. 

Second vegetative period 

The second vegetative period in the life cycle of the individual 
begins in September, when the roots protrude from the base of 
the primary bulb. These are few in number, three in most cases, 
and do not respond positively to gravitation. The root rudiments 
originate just below the region where the vascular strands from the 
dropper give off branches into the stem apex and the bulb scales. 
These stages are shown at r in figs. 24, 26, 27. The roots grow 
nearly straight from the point of origin to their limit of growth. 
They maintain a spatial relation toward each other so as to be 
equidistant in the soil. The oblique position of the base of the 
bulb makes the cluster of roots which radiate from the base appear 


In the 


tion is but slightly changed. The position of the roots in the soil 
is such as to define a nearly flat cone, with its axis in line with 
the axis of the apical dome. The angle taken by the axis of the 
cone varies slightly as the obliquity of the individual bulb differs, 
but lies close to 45 to the vertical. Under experimental tests, 
different positions of the bulbs used did not change the position 
of the axis of the cone of roots in reference to the axis of the apical 
dome, although the latter was itself placed in abnormal positions 
in respect to the vertical. Roots are rarely found within the coni- 
cal mass of soil outlined by most of the roots; when so found, 
the roots are very numerous and some are crowded away from the 
peripheral portion of the meristem cells in the base of the bulb. 




the bulb. In passing through the bulb the advancing tip of 




the root rudiment appears to come into contact with the cell walls 
at intervals only. This is indicated in sections by the presence 
of a cavity about the root tip, the bulb parenchyma being separated 
from the root itself by a space in which there appear no cell walls. 
In the cells at a little distance from the line of advance of the root, 
scattered starch grains appear, and the normal starch content 
remains in the bulb parenchyma farther from the root cluster 
(text fig. 2). It appears that the starch is first removed from the 
cells in the path of the advancing root rudiments, then as the root 
is organized and grows forward, the cells themselves are disinte- 
grated, leaving a clear pocket immediately about the conical end 

of the root. Only at the 
sides of these cavities were 

there any flattened cells, 
as if contact had occurred 
between the root tissue 
and the surrounding par- 
enchyma. But as the root 
issues from the base into 
the soil, there is a collar of 
cells about the base, which 
constricts the root at this 
point to. a slight degree. 

The number of roots in- 
creases with the age of the 
bulb, in large flowering individuals reaching 30 or 40. These all die 
at the close of the season, and a new set is formed for the next year, 
even if the new bulb is developed in situ. The roots have no con- 
tractile cells, and act only as absorbing organs; the shape of the 
bulb helps to anchor the plant in place. Root hairs are produced 
in abundance in a moist chamber, or in nature on those parts of a 
root which may pass through an air space in the soil. Where the 
soil is in close contact with the root, the root hairs are inconspicu- 
ous or undeveloped (in this connection see 8, p. 146). The men- 
stem tissue from which the roots arise continues to form new root 
rudiments for some time after the main supply of roots is formed. 
Thus a new crop of roots may be formed after the already developed 

Fig. 2. — Base of bulb, showing inception of 
roots in summer; X25. 


one has been cut off; or if in handling the bulbs the roots present 
become dry, new roots are protruded among the bases of the old. 
In sections young rudiments are to be found in the basal tissue 
until nearly the close of the growing season. 

The roots have a well-developed endodermis, formed of two, 
or rarely three, cell layers just outside the usually triarch stele 
(figs. 21, 22). The endodermis is delicate in the seedling, but well 
developed in the older roots. 


lished in the soil, and develops the bulb rudiment from the apical 
btid. There are two axillary buds; one of these is in the axil of 
the first leaf (cotyledon), the base of which elongated to form the 
dropper, and on the death of the seedling dried out to form the 
husk. The other bud is axillary to the inner scale, which forms 
the bulk of the bulb as it lies in the soil until the second growing 
season begins. In the primary bulbs neither of these buds ordi- 
narily develops further; but in older plants these, as well as the 
apical bud, form runners. During the mild periods of the winter 
and early spring the leaf is protruded from the bulb, and appears 
above the soil about the middle of March. The first leaves are 

cm. in 

than the older leaves, but in other respects like them. They last 
for six to eight weeks, and then disappear by retrogressive withering. 
During their activity the bulb rudiment at the base of the petiole 
enlarges to its full size as the secondary bulb. 

The starch stored in the primary bulb is used in part in the 
formation of the aerial structure (foliage leaf), in part also in the 
building of the new bulb. The bulb gradually disappears as the 
new bulb develops, the older tissues becoming free from starch and 
reacting to the Fehling test for sugar as the season advances and 
the new bulb enlarges. When the leaf begins to wither, in late 
May, there is practically nothing left of the primary bulb except 
its husk; this still incloses the base of the petiole, which forms the 
isk of the secondary bulb. After the death of the foliage leaf, 
e bulb is dormant until the roots are developed in September, 
as the first step in the third vegetative season. But in the interval 
the stem apdx organizes the foliage leaf, and organizes the buds 


350 BOTANICAL GAZETTE . [November 

in the axils of the two bulb scales as runner rudiments, in preparation 
for the next year. The details are similar to the same steps in the 
primary bulb. 

Immature stages 

Between the development of the secondary bulb and that of 
the bulb which bears a flowering shoot, an indefinite number of 
years may elapse. The least number of intervening seasons has 
been calculated to be three, making, with the first and second 
seasons, a minimum of five years from seed to seed. Many indi- 
viduals at any stage fail to produce runners in any one season, and 
their immature period is correspondingly lengthened. Others 
fail to gain any depth in the development of the new bulbs from 
runners, and these also lengthen the interval between the first and 
final bulbs. With the production of the flower the activity of the 
primary stem apex ceases, the subsequent bulbs being developed 
annually each from a bud at the base of the aerial shoot. In the 
immature individuals more than one bulb is formed in average 
seasons in four species, 2 more or less specialized structures being 
developed for this purpose. Most of the plants producing runners 
in E. americanum in any one season after the second bulb have 
three at a time. One of these is the apical bud, the other two are 
axillary buds. The relation of these three buds is shown in figs. 
28-33; these buds have been mentioned in a preceding paragraph 
as related to the development of the runners. The runners in the 
four species mentioned are not uniform in character, since in the 
development of the elongated structure of the runner different 
tissues are made use of by the plant in different species. Ery- 
thronium americanum is the most specialized in its development, 
but it is the clearest morphologically. 

Each of the runners in E. americanum comes from a bud within 
the bulb (text fig. 3), the buds being made up finally of two scale 
leaves and a stem apex inclosed by them. The buds are inserted 
upon the tissues of the bulb in such a manner that the outer surface 

2 These are E. americanum, E. albidum, E. propullans, and E. Hartwegii; others 
rarely multiply. In E. propullans the adult plants have a lateral runner in addition 
to the renewal bulb within the old. Under cultivation axillary buds in adult bulbs 
in most species develop into bulbs. 




of the external scale is united to the subtending portion of the 
bulb. As the buds begin their development, the first growth is 
eccentric and pushes the base of the bud outward from its first posi- 
tion, leaving the bottom of the bud free from contact with other 
tissues. Subsequent growth on the part of the bud is chiefly 
located in a zone about the base of the scale, close to the insertion 
of the inner scale upon the base of the bud. As the point at which 
the bud is united to the bulb tissue is above the zone of growth, 
the growing zone tends to push the base of the bud away from its 
original position and downward into the soil, as was the case in the 
beginning of the dropper. This elongation is confined almost 
entirely to the outer scale, the inner scale and the inclosed stem 


Fig. 3. — Space relations of buds in immature bulb of E. americanum: I, stem 
a Pex; 2, inner axillary bud; 3, outer axillary bud; Xio. 

apex being carried along passively, as the terminal bud in the 
runner (cf. fig. 13). The runner is thus formed by the elongation 
of the outer scale of the bulb rudiment, and in the case of the 
apical bud and the inner axillary one, they burst through the 
inclosing base of the petiole when the base of the bulb has been 
penetrated. In these runners the scale is definitely organized 
in the bud, before any elongation begins, and the base of the 
foliage leaf takes no share in the formation of the runner, as did 
tne base of the cotyledon in the seedling dropper. In other species 3 
the sheathing base of the foliage leaf contributes to the developing 

runner, as in the case of the cotyledon and dropper. The elonga- 
tion in this type of runner is the same as that in Tulipa (Robert- 

3 E. albidum for example. 


son 17, Irmisch 10) , in that the protrusion of the petiole base 
carries with it the bud inclosed by the petiole sheath. 

In E. albidum two runners are usual, while the species from the 

lv form but one. The 


length of the runner may be so much reduced as to pass unnoticed, 
but unusual conditions may demonstrate the presence of the 
typical structures not otherwise distinct. When but one runner 
is formed, or when the bulb is renewed in situ, the main stem apex 
is the active structure, the axillary buds if present remaining unde- 

There seems to be a moisture relation on the part of the growing 
runners. Plants of £. americanum growing in well-drained wood 
soil were found in several cases to have few runners; while other 
plants of similar general appearance, but in wet soil at the bottom 
of the same hill, showed runners in most individuals. It is notice- 
able about Baltimore that those individuals which grow in heavy, 
wet soils produce runners earlier and more abundantly than those 
of drier habitats. The development of abundant runners would 



of flowering plants in a colony of stated size. The conditions under 
which the plants grow are evidently directly related to flower 
production, as the abundance of bloom has been observed to be 
associated with abundance of moisture in each of the habitats 
examined during the work here discussed. 

In moist chamber experiments it was found that the tips of the 
runners were positively geotropic when lying on the top of saturated 
sphagnum, a little free water being present in the dish (fig. 23). 
In the absence of free water in the dish, the response was less 
marked, and in mere dampness it was a negligible quantity* 
testing to determine whether the downward growth of the tips 
under abundant moisture is due to a moisture response or to 
gravitation, experiments were made with moist sphagnum in 
different positions in reference to the growing runners. The slow 
growth and the sluggish response on the part of the runners made 
the experiments inconclusive. In the moist chamber a period ot 
two or three weeks was needed to carry through a set of tests. 




The slow growth of the runners made experiments with them 
unsatisfactory when using the centrifuge or klinostat, but the evi- 
dence obtained tends to confirm the indifference to gravitation noted 
under natural conditions. 

The lateral displacement of axillary buds, with little subse- 
quent elongation, occurs in species of Gagea (Irmisch io). In 
G. minima, G. lutea, and G. pratensis there are short protrusions of 
the bulb rudiment as if about to form a runner, but the elongation 
is not sufficient to burst through the base of the bulb. The sheath 
in which the new bulb is formed is the base of one of the leaves, 
which forms a pouch or pocket about the enlarging bulb rudiment. 
In Tulipa the base of the petiole not only forms the pocket imme- 
diately about the bulb rudiment, but elongates in such a manner 
as to carry the inclosed bud forward into the soil as the terminal 
bud in a hollow runner. This is the type mentioned above as 
occurring in Erythronium albidum and others in respect to the apical 
bud. In each of these cases the bud tends to become deeper than 
its predecessor, but in Allium vineale and A. Scorodophrasum 
(Irmisch io) the buds are borne upon independent bases, which 
elongate during the growth of the buds, so that the the buds are 
elevated from the first position and thrust out of the top of the bulb. 
In the cases first mentioned the buds are attached laterally to the 
bulb tissue from which they arise, and are thus unable to rise above 
their insertion, but must descend more or less sharply in the indi 1 
vidual species. 

Structure of runners 

The structure of the runner may be understood from the accom- 
panying diagram (text fig. 4). Such a bud as that of Allium 
vineale, which elongates in the region below the insertion of the 
bud scales, would be elevated above its original position; this is 
the normal erect bud. On horizontal rhizomes a bud assumes the 
position shown in 2 of the diagram at the close of the period of 
elongation. If this position were taken during active progress 
through the soil, a great amount of resistance would have to be 
overcome in pushing the erect bud forward. This exact arrange- 
m ent of parts has not been noticed in nature; but it occurs 
slightly modified in Erythronium propullans. In this species the 




lateral runner issues from the side of the shoot with the bud at 
the tip in an erect position. But the bud does not stand freely 
exposed above the general surface of the runner stalk. The region 
immediately behind the bud, where the scale and cauline tissue 
are fused, thickens vertically, so that the bud receives the support 
of the adjacent tissue for its whole height, and is therefore not 
subject to transverse strain tending to invert it upon its base. 

In the absence of supporting tissue for the height of the bud, 
the elongating runner would tend to advance the base of the bud 

Fig. 4.— Evolution of Erythronium type of runner: J, normal bud, terminating 
stem (Allium); 2-4, hypothetical stages, with horizontal stem; 5, anatropous bud 
of Erythronium runners; dotted lines and arrows indicate zone of elongation in each 


beyond the rest of the bud, as in no. 3, since the unsupported parts 
of the bud would be retarded by friction in passing through the 
soil. The tendency under such conditions would be for the adjacent 
surfaces of the bud scale and the stalk to fuse, as suggested in 
no. 4. Up to this point the scales of the bud have taken no part 
in the elongation of the runner or of the bud at the tip; but with 
the fusion of the bud with the surface of the runner stalk, either the 




I be 



runners of E. americanum the structure sketched in no. 5 is present. 
The zone of elongation is close to the organic base of the bud, 
while the bud scale is fused for its full height with the stalk of the 
runner. There are no disruptive strains in this case, for the 
external scale of the bud elongates as fast as does the cauline tissue 
of the runner; that is, the scale leaf, like a foliage leaf of the same 
plant (Robertson 17), elongates in the basal zone; the stem 
elongates in its apical region, and as the two are folded back upon 
each other, the two growth regions are continuous about the 
inclosed bud. The position of this region of elongation is indi- 
cated by the space between the diagonal dotted lines in no. 5. 

The cauline tissue in the runner is represented by the vascular 
bundles which unite the terminal bud in the runner tip to the base 
of the parent bulb. The presence of foliar tissue on the same side 
of the runner is indicated by the course of the vascular bundles, 
which turn backward at the base of the scale, and supply the inner 
face of the axial wall of the dropper in the same manner as the thin 

abaxial wall, where there is no tissue other than the scale. (Cf. 
text fig. 1 .) 

The runner habit 

The plumule (stem apex) in the cotyledonary sheath does not 
form any bud scales which might by their elongation produce the 
dropper; but the walls of the cotyledonary sheath elongate after 
the primary root is established, and develop the dropper with 
the plumule as the terminal bud therein. This appears to be 
constant for all the species of the genus. In the subsequent 
stages in the several western species, and in E. albidum and E. 
mesochoreim, the main stem apex is thrust out as the runner bud 
within the elongated sheath formed by the clasping petiole. This 
is exactly comparable to the dropper of the seedling, in that the 
bud is carried forward passively and contributes nothing to the 
development of the structure. In the case of the second runner of 
& albidum and of each of the three runners in E. americanum, the 
^nner is formed by the bud itself forming the runner sheath by 
the elongation of its outer scale. The development of this type 
0I " runner introduces a new structure into the series present in 
the life cycle of the other species, and the degree to which this 




innovation is developed may b6 taken as an index to the divergence 
from the original type. On this basis the most divergent species 
is E. americanum, with E. albidum and E. mesochoreum in order 
toward the type. Erythronium propullans is a special case, in 
which the bud axillary to the foliage leaf (text fig. 3, bud 2) in 
immature plants (as of E. americanum) is functional in the mature 
individuals, being elevated above the base of the bulb by the 
growth at the base of the aerial shoot, as this raises the base of the 
leaves to their final position. In floral characters it is most nearly 
related to jE. albidum. The reduced size of the flower is probably 

1 9 

Fig. 5.— Offshoot of £. propullans, showing structure and vascular elements in 
sections: A , C, on base of older off shoot B; b, bud of offshoot; c, cavity about bud. 

due in a measure to the interference with normal nutrition of the 
flower caused by the diversion of a portion of the vascular supply 
of the peduncle into the developing runner. The bud from which 
the lateral runner develops is inserted on the base of the peduncle 
in such a manner that in the early stages of growth as a runner 
the base of the peduncle elongates and thickens. As the clasping 
petioles (sk, text fig. 5) hold the upper part of the peduncle from 


bend. At this stage 

all the 

vascular bundles are dn 
base of the peduncle (A) 



elbow of the peduncle, certain of the peduncle bundles produce 
branches which grow with the advance of the bud and maintain 
connection with the main vascular supply of the plant (B) ; but in 
so doing the total supply available to the peduncle is divided 
between the runner and the peduncle itself, and the reduced size 
of the flower appears to be the result. 

In the species east of Colorado the aerial portions of the plant 
disintegrate with the ripening of the seeds, the fruit being prostrate 
when the seeds are ripe. The species of the Rockv 



ripen the stalk becomes a stiff and elastic wand, from the top of 
which the seeds are shaken. In these the capsule is revolute only 


among the first species the seeds are thrown out of the capsule by 
shaking; in the second group they are released by the rolling back 
of the inclosing walls of the prostrate capsule. The Eurasian 
species E. Dens-canis belongs to the second group in respect to its 
seed dispersal. 

The western species differ from the forms of the eastern states 
in details of bulb development. In the eastern forms the region 
of fusion between the bulb tissues and those of the bud is small; 
and the sheath inclosing the bud, either in the case of the runner 
bulbs or when renewed in situ, is free from the developing bulb 
for most of its surface; but in the western forms the elongated 
bulb has a long fusion zone along the axial side, where the base 
of the sheath and the surface of the inclosed bud scale are fused. 
This appears as a ridge, from the edges of which the husk, in 
fully formed bulbs, extends around the rest of the bud. The 
bulbs are as a rule much more attenuate in the western forms than 
m the eastern, both in the young and in the adult stages. In 
immature bulbs the tip of the new bulb is often not at all lower 
than the base of the preceding bulb ; this is due to the elongation 
upward on the part of the bud scales at a rate equal to that of the 



older bulbs the descent is decreased, but the elongation of the 
°ulb scales upward from their bases produces the long bulbs 
characteristic of the group. In the bulbs examined, the species 



from the western states have thinner husks than those of the 
eastern states, except E. Hartwegii, which has a tough and thick 
one like that of Tulipa. This form grows in an adobe soil, subject 
to high temperatures and considerable pressure when the soil 
dries, which may have its influence in the development of the 
heavier husk (Warming 22). The Eurasian species (E. Dens- 
canis) has a very delicate husk, which becomes fragmentary during 
the development of the new bulb, so that the bulb is but slightly 
protected. The development of the bulb in this species is similar 
to that of the western forms, but the zone of fusion on the axial 
side of the bulb rudiment is more nearlv the full height of the 

in these, extending almost 


the scales are fused together along their adaxial surfaces and the 

much farther from 

than is the case in any other species. The base of the petiole 
becomes charged with starch, and indistinguishable in appearance 
or function from a bulb scale. This explains the absence of a 
husk, formed in the other species from the drying of the sheathing 
petiolar sheath. The seedling of this species is similar to that 



United States. It is probable that the period of blooming is 
attained even more slowly by E. Dens-canis than in the other 
species, if the same relation of depth and flower production is 
maintained. In some of the western species grown in the Botanical 
Garden of Johns Hopkins University, it was noticed that they 
bloomed when the bulbs were much smaller in proportion to the 

ge mature 

It may 


and in the western species, and that the bulbs continue to descend 
gradually during a long period after the blossoming habit is inaugu- 
rated. Such a point is physiological rather than morphological, 
and will not be discussed further at this time. 

In the runners of E. americanum occasional "stimulation 
growths" have been noticed, apparently related to sudden exces 

of water. 



through the tip of the runner, just as the runners burst the petiole 
sheath when first issuing from the bulb. The next scale of the 
bulb rudiment forms the runner extension, and the stem apex 
with a small scale is carried forward as if under normal conditions. 
The necessary nourishment for the added growth comes from the 
parent plant, through the stalk of the runner to the point where 
the terminal bud was inserted. Then the vascular bundles of the 
bud, following down through the bud scale, convey material to 
the stem apex in the tip of the extension of the runner. In two 
cases, in the spring of 1908, this extension of the runner by the 
protrusion of the terminal bud through the end of the runner 
sheath had been repeated a second time. 

Development of the mature plants 

The production of the flowers, which mark the maturity of the 
individual plant and the termination of the cycle of stages from 

seed to seed ("text fipr fiV i« fl.Qsnr.ia.teH with several changes from 




dition. During the whole period from seedling to this stage, the 
primary stem apex has persisted, and from it have come the suc- 
cessive leaf rudiments which developed into either bulb scales or 
foliage leaves as the conditions demanded. The accompanying 
tabular view of the activity of the primary stem apex will show 
the general relation of these leaf rudiments. In addition to the 
outgrowths from the apical dome in the form of leaf rudiments, 
there have been various buds, axillary to the inner scale, which 

apex from time to time. The leaf 
rudiments developed in these axillary buds are not considered 
in the tabulated series of products from the stem apex. The bud 
in the axil of the outer scale does not arise from the apical tissue 
directly, but from the mass of meristem at the base of the scales. 

The final structure developed from the primary stem apex, 
after passing through the immature stages, is the flower bud of 
the mature individual. In the immature stages the apical dome 
persists at the base of the bulb, producing successively the several 
bulb scales and foliage leaves. In the mature bulb the apical 
dome becomes surrounded by the foliage leaf rudiments (fig. 25), 




and the whole structure becomes elevated above its original 
position by the elongation of the tissue below the insertion of the 
leaves. In the production of the flower bud, at the summit of 
the aerial shoot, the original stem apex comes to an end. The 
renewal bud for the next bulb is formed as a bud in the axil of 
the inner bulb scale, at the base of the shoot (figs. 256, 26, 27). 








Foliar structures 





First leaf 

Second leaf 
Third leaf 

Fourth leaf 

Fifth leaf 
Sixth leaf 

Seventh leaf 

Eighth leaf 
Ninth leaf 

Tenth leaf 

Eleventh leaf 
Twelfth leaf 

Thirteenth leaf 

Fourteenth leaf 
Fifteenth leaf 

Sixteenth leaf 


First scale 
Second scale 

First green leaf 

Third scale 
Fourth scale 

Second green 

Fifth scale 
Sixth scale 

Third green 
• leaf 

Seventh scale ■ 
Eighth scale 

Fourth green 

Ninth scale 
Tenth scale 

Fifth green leaf 

(and flow 















er stalk)f 


Bulb scale 


Outer scale 
Inner scale 


Bulb scale 


Bulb scale 


Outer scale 
Inner scale 






* If the bulb is renewed in situ, the base of the leaf becomes the husk of the new bulb; if the new 
bulb develops as a runner bulb, the runner sheath forms the husk. 

t With the development of the floral axis the activity of the primary stem apex ceases, a branch 
apex forming the renewal bulb thereafter.. 


This bud is evidently the homologue of bud 2, of the immature 
stages, as indicated in- text fig. 3 . The bud is at first distinctly 
in the axil of the scale, but with the upward growth of the adjacent 
shoot it is lifted from its position to one on the base of the shoot 
itself (figs. 27 and 29, 3). 

The apical dome of this bud develops bulb scales during the 




summer and develops into the full-sized bulb during the active 
period of the aerial shoot at the base of which it lies. Starch for 
storage is derived both from the photosynthesis of the leaves 
and from the deposits in the old bulb within which the new bulb 

June Dec 


mchetS deep 

Fig. 6.— Stages in life cycle of E. americanum: under natural conditions the third 
bulb may repeat itself indefinitely, gaining little in depth, being renewed in situ for 
one °r more seasons. 

one °r more seasons, 

is formed. 



°* the next flower. This process is repeated indefinitely, the 

362 BOTANICAL GAZETTE ' [November 

individual becoming more robust with time, but otherwise remain- 
ing the same. A very slight increase in depth occurs as time 
passes, since the new bulbs are a little larger than their predeces- 
sors, and the new bulb is formed below the center of the old one 
which it replaces. But a bulb once flowering may revert to the 
sterile condition after a series of years. Bulbs which have been 
dug from the soil and allowed to become somewhat dried out 
are apt to "break," that is, to produce two or three small bulbs 
instead of the single large one customary in the flowering plants. 
Bulbs so produced are sterile for at least two years, as found by 
experiment. This is especially apt to occur if the bulbs are dug 
while the shoot is actively growing upward through the soil in the 
spring. Removal of the soil close to a bulb of a flowering plant 
often causes such to revert to the immature condition and produce 
runners, even though the flowering of the current season is not 
hindered by the changed soil conditions. 

Individual plants of any age may force the bulb rudiments 
into abnormal development under the stimulation of unfavorable 
conditions. A large number of bulbs of various sizes were dug 
soon after the leaves had pushed through the soil. These with 
a little soil were left in a tin box for several days; the box was 
well wrapped and left in a cool room in the meanwhile. Upon 
inspection ten days later, the leaves were found to have shriveled 
away, but the bulbs had formed a small bulb from each of the buds 
which under normal conditions would have developed runners. The 
starch present in the old bulb and in the leaf furnished the material 
for the new bulbs, aside from water, which was in the moist soil. 
This was repeated several times, the exact details varying with 
the exact condition of the bulbs when dug from the soil. Thus if 
the runner is already pushing through the base of the bulb, or 
is further developed, it continues to elongate for several days, but 
in the end will form the terminal bud into the bulb; it apparently 
transfers to it the substance of the old bulb and runner stalk as 11 
under normal conditions. 

In the descriptions of the liliaceous ovary, the placentae are 
usually said to come from the incurved margins of the carpellary 
leaves. In Erytkronium, however, the margins of the carpels 




appear to come into blunt contact, but do not curve inward. The 
margins unite along the line of contact, and the partitions are formed 
from a median rib projecting inward from each carpel (figs. 34, 35), 
meeting at the center of the ovary (Temple 21). In the full-grown 
ovary, as at fertilization, the evidence for this interpretation of 
the case is found in the double bundles along the line of dehiscence 
in the walls of the ovary, and in the 
presence of a layer of cells rich in pro- 
toplasm which passes around the end of 
the abutting partitions as they come 
into contact at the center of the ovary 
(text fig. 7). The densely protoplasmic 
cells which line the ovary along the line 
of the placentae continue around the 
end of each median wing, as shown in 
the photograph. This series of surfaces 
of slight contact determines the dehis- 
cence of the ripe capsule. 

Occasional plants are found with 
sterile anthers on one or more of the 
stamens, some patches showing many of 
the plants with no functional pollen. It 
has been found that the sterility of the 
anthers begins at least as early as the 

divisions of the pollen mother cells, as , FlG ;.. 7 -~ Transve r rs ^ secd ° n 

buds have been examined in which three 
°f the anthers were normal, and had 

of fertilized ovary of E. ameri- 
canum, showing abutting 
placentae (/»), two-celled em- 

normal pollen grains; while the sporo- bryo («), and double bundle 
genous tissue of the others was degen- at ed * es of fused carpe,s (b)i 


crating (fig. 35, w> w ') ; the other 
tissues of the anthers were normal in appearance. In the field it 
has been noticed that the plants with dark pollen on the stigma 
had the larger fruits and the larger number of seeds. Plants 
having pale anthers often lacked pollen entirely, and their seed- 
developing power seemed to be deficient also. The plants of the 
two types, fertile with dark stamens and poor with pale anthers, 
each occur in patches often of considerable numbers. This may 


account for the poor set of seed in such cases, for insects working 
over the patch would not bring in so much viable pollen to the 


Development of seeds 

The fertilized ovule enlarges rapidly, mainly through the growth 
of the embryo sac. The sac destroys the nucellus except a crushed 
remnant at the bottom of the sac. The raphe and chalazal spur 


The endosperm develops slowly, remaining for a considerable 
time as a layer lining the wall of the sac between the base and the 
suspensor at the tip. The embryo first develops a considerable 
mass of cells in the micropylar end of the sac, from the inner free 
surface of which the functional embryogenic cells are developed 
(Coulter 7). This inner surface may become lobed, and then 
gives the condition called polyembryony by Jeffrey (13) and 
Schaffner (20). Twin embryos developed in one case out of 
several hundred seeds germinated in this work; these were united 
near the upper end, but completely free below, and their vascular 
systems were distinct. These embryos probably were formed 
from two adjacent lobes from the free surface of the suspensor, 
becoming united through the incorporation of some cells in common 
from the base of the lobes. 

In pushing into the soil, the elongation 


distinct in each of the two embryos, producing the condition shown 

(fig- 36). 

The general conditions of the sac, endosperm, and suspensor 


studied by Modilewsky (ik). In both cases there is a thin 

lining of free endosperm nuclei along the walls of the sac, a con- 
siderable suspensor at the tip, and a mass of deeply staining cells 

the base of the sac. In Erythronium these cells form a 
erable mass, and extend backward to the termination of 


the raphal bundle at the chalaza. These cells stain deeply even 
in ripe seeds, and the adjacent cells of the remnant of the nucellus 
remain in close contact with the center of the base of the endosperm 
in the ripe seed. This would indicate the purpose of these cells 




to be that of transfer agents for substances from the chalaza 

to the embryo sac, and thence to the forming embryo and 

When the seed reaches its full development the embryo sac 
is filled with hard endosperm of reserve cellulose. The cells in 
the endosperm have their longer axes directed inward from the 
periphery, curving somewhat toward the micropylar end of the seed. 
Through the center of the endosperm there is sometimes the same 
cellular endosperm as elsewhere, more frequently this is nearly solid 
cellulose. Immediately about the embryo there is usually a small 
mass of less firm endosperm, which in some cases extends to the sur- 
face of the seed, filling the space earlier occupied by the suspensor. 
It is probable that germination is aided materially in such cases, as 
moisture can more promptly reach the embryo by this path than 
would be the case if only thick-walled endosperm w T ere present. 
At the base of the seed the endosperm shows considerable shrinking, 
with the remnant of the nucellus remaining in contact with the 
endosperm for a small area near the center. This secures contact 
between the hard endosperm and the spongy spur, and even when 
the spur has disappeared the inner layers of the tissue at the chalaza 
remain in intimate contact with the base of the endosperm. As 
the endosperm along the axis of the embryo sac is the last to be 
deposited, it is probable that the mass of spongy cells in contact 
with the base is of advantage in the imbibition of water preliminary 
to germination. In different seeds the exact appearance of the 
central portion of the endosperm varies from nearly solid, with 
infrequent cell cavities, to a condition similar to that of the pe- 
ripheral endosperm; the latter is less common than is the solid 
core mass. The exact distribution of the cell cavities in the endo- 
sperm is probably in close relation to the moisture conditions 
during the late stages of ripening of the seed. In germination the 
embryo first enlarges to occupy the space filled by the spongy 


in which it lies; then the cotyledon, acting as a 

haustorial organ, dissolves a path for itself along the axis of the 
endosperm. The solvent action extends along the lines of cell 
cavities toward the periphery, and the endosperm is absorbed almost 


Completion of the cycle 

The individual which began with the germination of the seed 
and the organization of the stem apex in the sheath at the base of 
the cotyledon, has been followed through the series of changes 
normal to its development as Erythronium americanum, and the 
variations have been indicated between this species and the others 
of the genus. The cycle of the sporophytic generation is thus 
completed, and the original stem apex culminates in the formation 
of the flower parts of the first blossom produced by the bulb on 
reaching maturity. It was shown that the species were structurally 
related with a large group, the western species, behaving similarly, 
the others diverging more or less from the habits of these forms. 
The structures involved in the divergence from type are those 
most active in the immature stages, and most freely produced in 
E. amcricannm. This species, therefore, has been regarded here 
as the most remote from the ancestral form, since the majority 
of the species are of the more simple type of development in the 
vegetative habits. The species having the uniform vegetative 
habits are native to the region from the Rocky Mountains west- 
ward, becoming more abundant in species as the Pacific coast is 
approached, and culminating in a series of habitats in southwestern 
Oregon. This may be assumed to be near the original home of 
the genus, from which the distribution and differentiation into 
the present habitats and species have occurred. It is probable 
that the lines of migration have followed the present lines of 
specific distribution, especially in the United States. The single 
species of Eurasia combines the bulb characters of the western 
species with the withering aerial parts of the eastern forms. The 
character of the habitats in which this species occurs may be so 
nearly uniform that no marked variations have been developed from 
the original type, beyond that involved in the prostrate fruit. Ants 
have been observed carrying the seeds of E. americanum, and it is 
probable that they aid in distribution in each of the species having 
seeds with fleshy raphe or spur tissue. This would include E. Dens- 
cams as one of the myrmechochorous species, as a considerab e 
chalazal spur is present, comparable to that of Viola or Sanguinary 
(Beal i), but not so large as that of E. americanum orE. albidum. 



The points presented in this paper 


follows : 

The undifferentiated embryo begins to elongate in the fall and 
organizes a rudimentary stem apex in a narrow cavity at the base 
of the cotyledon. In germination the radicle is thrust into the 
soil during the winter, the stem apex following immediately behind 
the base of the radicle, in the cavity mentioned. The hypocotyl 
is represented by the fusion region between the vascular supply to 
the stem apex and the main vascular system of the seedling; it 
takes no part in the development of the seedling. After absorbing 
the endosperm, the cotyledon is elevated into the air, and acts 
as the first photosynthetic organ. Either two or three vascular 


es are present, varying with the species. 

During the activity of the cotyledon as a leaf, the cavity "in the 
base of the cotyledon about the stem apex elongates to form a 
slender sheath, with the stem apex inclosed as a terminal bud at 
the tip. The " dropper" so formed is positively geotropic in 
response, as is the primary root: the runners and roots of later 
stages are not positively geotropic. At the close of the season 
the terminal bud of the dropper is isolated in the soil by the wither- 
ing of other parts of the plant. 

During the summer the first foliage leaf is organized by the 
stem apex and becomes functional the following spring. Roots are 
Protruded at the base of the bulb in the fall, which marks the begin- 
nm g of the second vegetative period. This sequence of develop- 
ment is repeated in the subsequent seasons to the mature bulbs. 

The stem apex becomes inclosed by the base of the foliage leaf, 
and there forms a bud from which the next bulb will develop. 
When the bulb is renewed in situ, only this bud develops; this 
ls also the rule in the species producing but one runner. In E. 
tinericanum two buds are usually formed in addition to the main 
bu d, each of which elongates as a runner and forms a bulb. These 
additional runners spring from buds axillary to the leaf, or to a 
b ulb scale. When but one runner is developed normally by a 
s Pedes, it is formed from the elongation of the base of the petiole, 
w hich carries the bulb rudiment into the soil as a terminal bud, 


as in Tulip a. The additional runners when present, and all three 
in £. americanum, are developed from the elongation of the outer 
scale of the bud, the inner scale and the apical dome forming the 
terminal bud of such a runner. 

The vascular connection between the stem apex and the point 
of insertion of the runner is to be considered cauline, the rest of 
the runner tissue as foliar and apical. The structure of the runner 



of its external scale being fused with the upper surface of the support- 
ing stem; the latter maintains vascular connection between the 
base and the apex of the structure developed. The zone of growth 
of the fusion structure is located as in normal rhizomes, close to the 
tip of the structure; both the scale and the cauline tissue take 
part in the elongation. 

mature plants form the floral shoot from 





ment. The bud in the axil of the outer scale usually dies without 

habit as the basis 

divided into two groups, one occurring typically at low elevation, 
and producing two or three slender runners from short bulbs, the 
other at higher elevation and forming but one runner and develop- 
ing into slender bulbs. A form intermediate between the two 
occurs in western Kansas and Nebraska. An aberrant form occurs 
in a restricted locality in Minnesota, having a lateral runner 
developed from a bud in the axil of a foliage leaf. 

The interval between germination and flowering on the part ot 
an individual is at least six years. This minimum is liable to in- 
definite increase under average conditions, by the interpolation o 
additional immature bulbs because of the formation of the runner 
bulbs no deeper than the parent bulb, or by the suppression of 
runners in a particular year. 

The distribution of the species points to the Pacific coast as 
the probable home of the genus, the present distribution being 
the result of migration along lines connecting the habitats ol t 



present forms. The species are found to vary from the type found 

in the assumed original habitat approximately in proportion to the 

distance of migration. The species of Eurasia combines features 

of the two groups in its vegetative characters. 

The roots radiate from a point near the base of the bulb, but 

show no definite response to gravity; they maintain a definite 

space relation to each other, defining the surface of a cone in the 

soil, the axis of which is about 45 to the vertical. In development, 

the roots arise in September from cells just below the vascular 

base of the bulb, and grow in straight lines during their elongation. 

In penetrating the bulb tissue, the cells are apparently dissolved 

by the root tissue, making a cavity about the advancing root 

rudiments. The number of roots increases with the age of the 


From the foregoing study the following conclusions may be 
drawn: The delayed development of the embryo is associated with 
a large store of endosperm; this is drawn upon by the germinating 
embryo during the season when vegetative activity is low; the 
young seedling is established in the soil early in the spring, the 
endosperm furnishing the needed materials for its development. 
With the exhaustion of the reserve material of the seed, the primary 
root is developed, and the cotyledon is elevated into the air and 
light; the cotyledon is the only leaf exposed to the light by the 
seedling. The stem apex, located in a narrow cavity in the base 
of the cotyledon, is carried forward by the elongation of the embryo, 
and, after the elevation of the cotyledon, is carried farther into the 
soil by the elongation of the walls of the cavity. The short period 
°f vegetative activity, and the prompt descent of the stem apex in 
the dropper, would indicate adjustment to short growing seasons; 
the brevity of the active season is a feature of the life cycle. The 



« — i — 

"i some cases these are strictly comparable to the dropper of the 
seedling in structure, in others they involve new developments; but 
the result is the same in either case. 

The persistence of the original stem apex until the establish- 


merit of the flower axis allows for the repetition of an indefinite 
series of immature bulbs, formed from runners or in situ, but 
introducing no new structures during the whole period of immature 
development. With the formation of the flowering shoot, the 
vegetative structures become secondary in importance, and the 
renewal bulb is developed from an axillary bud at the base of the 
shoot. The continuation of the individual is thereafter without 
vegetative multiplication normally, the increase being secured by 
the seeds. 

The geographical distribution of species and their relation to each 
other in structural details indicate that the genus originated in 
that region of the Pacific coast now included in the state of Oregon, 
and has been distributed along lines approximately following the 
present habitats of the several species. In the progress of migra- 
tion the advancing species developed special methods for rapid 

forms has become 

numerical increase. 

In the development of means of vegetative multiplication 



the seedling the base of the cotyledon, in the western forms the base 
of the petiole) was followed by the elongation of the scales of 
axillary buds, thus forming additional descending axes, each of 


species has developed a runner from a bud axillary to the foliage 
leaf, apparently being derived from the adjacent form in which 
the second runner arises from a bud axillary to the inner scale. 
The production of this lateral runner is confined to the flowering 
plants, since only in these is the leaf axil elevated above the base 
of the bulb. This form is very restricted, and appears to be the 
species most recently derived from the parent stock, or from some 
other species as these are now known. 

The general development in the genus would confirm the 
assumption that it is related to Tulipa, especially through ?• 

Johns Hopkins University 

Baltimore, Md. 




2. Blodgett, Frederick H., Vegetative reproduction and mult 

in Erythronium. Bull. Torr. Bot. Club 27:305-315. 1900. 
3- -, The stem offshoot in Erythronium propullans Gray. John 

June 1900. 


mum. Mem. Soc. Nat. et Math. Cherbourg 30:1896-1897. 

5. Campbell, D. H., Elements of structural and systematic botany, pp. 
146-152. Boston. 1890. 

6. Chrysler, M. A., The development of the central cylinder in Araceae 
and Liliaceae. Bot. Gazette 38:161-184. 1904. 

7« Coulter, John M., Contribution to the life history of Liliiim Philadelphia 
cum. I. The embryo sac and associated structures. Bot. Gazette 
23:412-452. 1897. 

8. Frienfelt, S., Studien uber die Wurzelen krautigen Pflanzen. Flora 
91:115-208. 1902. 

9. Hofmeister, W., Neue Beitrage zur Kenntniss der Embryobildung der 
Phanerogamen. II. Monocotyledonen. 1861. 

10. Irmisch, T., Zur Morphologie der monokotylischen Knollen- und Zwei- 

belgewachse. Berlin. 1850. 
11 • , Beitrage zur vergleichende Morphologie der Pflanzen. Bot. 


Zeit. 21 : 137-142. 1863. (Tulipa) 

Nat. Gesell. Halle 7:175-226. 1863. (Erythronium) 

Morphologie der Pflanzen. Abhandl 

2 3- Jeffrey, E. C, Polyembryony in Erythronium americanum. Annals of 

Botany 9:537-541. 1895. 
H> Jost, L., Plant physiology. English translation. 1907. 

Y, J., Zur Samenentwicklung einiger Urticifloren. Flora 98: 

x "50- 1908. 

l6# Ri ^bach, A., Physiological observations on some perennial herbs. Bot 

^*aiis 30:171-188. 1900. 
x 7. Robertson, Agnes, The droppers of Tulipa and Erythronium. Annals 

of Botany 20:423-440. 1906. 
I8 - Sachs, J., Ueber die Keimung von Allium Cepa. Bot. Zeit. 21:37-62. 

*9. Sargant, Ethel, A theory of the origin of monocotyledons founded on 

the structure of their seedlings. Annals of Botany 17-1-92. x 9°3- 

history and cytology of 

Erythronium. Bot. Gazette 31:369-387. 1901. 
Temple, Charles E.. The structure of lilv pi 

Science N.S. 29: 




009. p. 124 




All figures drawn on the same scale, which is indicated by the measured 
line; drawings made by means of a vertical projecting lantern from the stained 

Fig. i. — Dormant embryo in endosperm; *, remnant of inner integument. 

Fig. 2. — Embryo about December i, still within seed; s, position of stem 
apex; e, endosperm. 
* Fig. 3. — Descending tip of seedling; stem apex in slit at a. 

Fig. 3a. — Cross-section of seedling at line a-boi fig. 3. 

Fig. 4 — Tip of cotyledon in longitudinal section, showing haustorial cells 
(h) and vascular cells (v). 

Fig. 5. — Cross-section of cotyledon; v, vascular bundles; s, stomata. 

Figs. 6-13.— Stages in the development of the dropper about the stem 
apex (s). 

Figs, ii, 12. 

and stem apex (s) ; t, plumule trace. 


Fig. 13. — Stem apex as terminal bud in tip of active dropper, as in April. 



Magnification the same as in plate VIII, except in figs. 21, 22, 23, 25, in 
which the magnification is indicated. 

Fig. 14. — Stem apex in late June at bottom of bulb. 

Fig. 15. — Stem apex (s) and rudiment of foliage leaf (/) in July. 

Figs. 16-20. — Cross-sections of young dropper, from base of cotyledon 
downward, showing hypocotyl region (17), separation of root and dropper 
(18), and dome of stem apex (20). 

Figs. 21, 22.— Cross-sections of active and dead root steles; e, endodermis; 

Fig. 23.— Active runner marked to show zone of maximum elongation after 

three weeks' growth; natural size. 

Fig. 24.— Base of bulb, showing sheathing petiole (p), stem apex (*), and 
root rudiment (r), as found August 1. 

Fig. 25.— Outline of stem apex in flowering bulb, showing flower rudiment 
(/) inclosed by the foliage leaves (/) and the location of the renewal bud (b) ; 


Fig. 26.— Renewal bud from the same section showing apical dome above 
root rudiment below the scale; July 

Fig. 27. — The same structures about August 1, on the same scale. 

plate x 






\. I 

■ J «ttl V, 1 )Q 








0.1 mm 0.1** « 

i —J 




A ' 

m 1 

HlJODOtTT on AM TffffOA 'If ! V 





Fig. 28. — The three buds in the immature bulbs of E. americanum; 
main stem apex; 2, bud axillary to leaf; 3, bud axillary to scale; 1 and 3 
on the same section. 

Fig. 28a. — Sagittal section of bud 3, to show apical dome. 

Fig. 29. — Outline of entire section shown in fig. 28; relation of buds 1 
and 3. 

Fig. 30. — Similar section in reference to bud 2; in each the sheathing base • 
of the petiole is p. 

Figs. 31-33. — Cross-sections of bulb rudiment, showing relation of the same 
structures as shown in figs. 28-30; the sections follow in order of their numbers 
but are not consecutive. 

Figs. 34, 35. — Cross-sections of developing flower bud in early September 
and early December, on the same scale and in the same orientation; /, foliage 
leaves; p, petals; s, stamens; 0, ovary wall; /, line of dehiscence (lateral margin 
of carpel); w, wing-placenta from median region of carpel; m, pi f , normal and 
degenerating microspores. 

Fig. 36. — Twin embryos of E. americanum, X5; the union of the two 
cotyledons shown below in outline. 




Irving W. Bailey 


In studying the phylogeny of plants there are certain principles 
or canons of comparative anatomy which have been formulated 
within recent years by morphologists and anatomists. Thus the 
application, to seedling plants, of Haeckel's law of recapitulation 
for stages in ontogenesis has been strikingly illustrated by Stras- 
burger, Goebel, Jeffrey, Eames, and others. Furthermore, 
the persistency of ancestral characters in certain regions of plants 
has been well established. In this connection the researches of 
Solms-Laubach, Scott, and Jeffrey have shown conclusively 






ill. Jeffrey has 

the importance of 


recapitulation of ancestral conditions. The importance of hyper- 
trophied or wounded areas as the seat of reversion to primitive 
characters is strongly appreciated by zoologists. This principle 
has also been applied to the traumatic areas of plants by Jeffrey, 
who has pointed out the traumatic reversionary origin of resin 
canals in the wood of the higher Abietineae, 3 certain Sequoiineae, 4 
and the older Araucarineae, 5 which normally possess none of these 

Contributions from the Phanerogamic Laboratories of Harvard University, 
no. 24. 

3 The comparative anatomy and phylogeny of the Coniferales. I. The genus 
Sequoia. Mem. Boston Soc. Nat. Hist. 5:441-459. pis. 68-71. i9°3- 

* The comparative anatomy and phylogeny of the Coniferales. II. The Abietin- 
eae. Mem. Boston Soc. Nat. Hist. 6:1-37. pis. 1-7. 1904. 

* The comparative anatomy and phylogeny of the Coniferales. I. The genus 
Sequoia. Mem Boston Soc. Nat. Hist. 5:441-459. ph. 68-71. i9°3- 

sThe wound reactions of Brachyphyllum. Annals of Botany 20:383-394- P Sm 
27, 28. 1906. 

Araucariopitys, a new genus of araucarians. Bot. Gazette 44 : 435 
28-30. 1907. 

Botanical Gazette, vol. 50] 




structures; and has demonstrated the presence of traumatic mar- 
ginal tracheids in the wounded wood of Cunninghamia sinensis. 6 

An interesting parallel to the work of Jeffrey upon traumatic 
reversions in the Coniferales has been noted by the writer in the 
traumatic reversions of wounded oak wood. Owing to the con- 
troversy which exists as to the relative primitiveness of the Abietin- 
eae and Cupressineae this occurrence of traumatic reversions in 
a dicotyledonous genus is of particular interest in demonstrating 

the application of the principles of experimental morphology to 

Before describing the reversionary characters of oaks, it will 
be well to have clearly in mind the normal structure of the wood 
of existing oaks, and also that of ancestral types. As is well 
known, the secondary xylem of living oaks consists of vessels, 
fibers, tracheids, and parenchyma. The last occurs vertically 
as wood parenchyma, and horizontally disposed in plates of tissue 
extending radially, the so-called primary and secondary medullary 
rays. The "primary rays" (fig. i) are the distinctive feature of 
oak wood, the "silver grain," and are broad, fusiform masses, 
many cells in width, which are supposed to originate as inclusions 
of fundamental tissue between the primary fibrovascular bundles. 
The "secondary rays" are thin sheets of tissue, and consist of a 
single row of cells when seen in tangential or transverse section 
(fig- i). These rays, unlike the 


originate only with secondary growth. 

In contrast to this type of structure occurring in the mature 
wood of extant oaks, Eames has shown 7 that certain miocene 
oaks do not possess large rays composed of homogeneous masses 
of ray parenchyma, but have in their place bands of aggregated 
smaller rays which are separated by fibers and wood parenchyma. 
These rays are homologous with the "false rays" of the Betulaceae, 
and lead to the conclusion that the so-called primary rays of extant 
oaks have been built up by an aggregation and fusion of numerous 



Pi- J/. 1908. 

Annals of Botany 22 : 59 3 

7 On the origin of the broad ray in Quercus. Bot. Gazette 49:161-166. pis. 
*> P. 1910. 


The writer has shown, 8 by . a study of numerous species of 


process exists in the rays of many living oaks, alders, birches, 
and hornbeams. Thus many American live oaks (fig. 5) possess 
bands of aggregated rays which are similar to the false rays of the 
Betulaceae. Among alders and birches species may be found 
with non-aggregated uniseriate rays, aggregated small rays, and 


In fact, in the genus Alnus species may be found which form a 
perfect series of transitional steps between alders with non-aggre- 

rhombifolia Nutt 



Further, Eames has shown that in the development of seedling 

oaks a similar series of stages occur. 
alba L. and Q. rubra L., which in t 


formed wood resembles 

chestnut in possessing non-aggregated uniseriate rays. In the 
further growth of the young plant, aggregations of rays develop, 
which by subsequent fusion constitute the large rays of the mature 




tures in living forms, show conclusively that the wood of primitive 
( Fagaceae and Betulaceae was characterized by the entire absence 
of large medullary rays. With the development of unequal 
seasonal temperatures, a highly organized storage system for foods 
became advantageous to plants, and the large rays of modern 
oaks have been evolved by an aggregation and fusion of numerous 
uniseriate rays to meet this demand. 

With these preliminary statements on the normal and ancestral 
features of oak wood, we may now turn to a more detailed consider- 
ation of the abnormal structures which exist in traumatic oaks. 
It has been pointed out above that traumatic regions are often 
the seat of "ancestral characters and show a series of stages similar 


the dicotyledons. Annals of Botany. Ined. 


to embryonic or seedling stages. We should accordingly expect to 
find in wounded oak wood a reversion to primitive ray structure. 
That this reversion does occur has been shown conclusively by 
the examination of numerous areas of wounded oak wood. In 
fact, in all cases a complete series of transitional stages in the 


development of compound rays occurred, 
of steps found by Eames in the development of seedling plants. 
The wood formed immediately after wounding possesses char- 
acteristically only uniseriate rays. In following layers aggrega- 
tions of the rays develop, with subsequent enlargement of the uni- 


or compound rays. During this com 


into ray parenchyma. 

In all cases, in securing material, burls and distorted tissues 
were carefully avoided and straight-grained traumatic tissue 

In fig. 



wood of Quercus nigra L. The so-called primary and secondary 
rays are characteristically developed; the former consists of a 

In contrast to this the 


traumatic wood of the same species may be seen in fig. 2. The 




United States. As may be seen, in this 



ous included fibers and wood parenchyma cells, evidences of fusion, 
are usually present. Fig. 4 illustrates the reversion of the wood 
in the vicinity of the wound to the non-aggregated uniseriate 

In figs. 5 and 6 are shown sections of the normal and the trau- 

Quercus densiflora Hook 


cess of compounding is clearly shown. The wounded wood of 

more primitive 



Figs. 7-10 illustrate the normal and traumatic wood of the well 
known Quercus alba, a deciduous oak with highly specialized ray 
structure. Figs. 8 and 10, tangential and transverse sections respec- 
tively of the traumatic wood, show that this species also reverts 
to ancestral type of ray structures. 

This reversion to ancestral conditions has been found by the 
writer in woody tissues subsequent to severe injury. Slight 
injuries, on the other hand, particularly in oaks with highly special- 
ized ray structure, often produce no morphological effect upon the 
wood. In a limited number of cases very slight wounds have 
produced a stimulation of the compounding tendency or accelera- 
tion of development rather than a reversion to ancestral structures. 
In fig. n may be seen a cross-section of Alnus sp. The lower 
portion of the figure shows none of the so-called false rays, but 
in the upper half numerous rays have developed abruptly outside 
a zone indicating slight injury. Fig. 12 shows a similar phenome- 
non in the young stem of Quercus densiflora. In the lower half 
may be seen normal seedling wood in which only uniseriate rays 
occur. Starting from a zone of slight injury in the middle of the 
section, a compound ray has suddenly appeared in the upper 
portion of the figure. From this we see that very slight injuries 
produce an acceleration in the formation of compound rays. 


1. The phylogenetic importance of traumatic areas as the seat 
of reversion to primitive structures is well illustrated by the speci- 
mens of wounded oak wood which have been examined by the 

2. In traumatic wood progressive stages are found which are 
similar to the stages of recapitulation found in the seedling, and 
equivalents to the condition found in adult miocene oaks. 

3. Woody tissues in the immediate vicinity of a severe wound 
show only non-aggregated uniseriate or small rays. In subse- 
quently formed tissues the gradual building up of the compound 
ray may be traced in a consecutive series of steps to the normal 
homogeneous large ray of the adult wood. 

4. On the basis of traumatic and developmental as well as paleo- 


botanical evidence, the large homogeneous masses of ray paren- 
chyma, or the so-called primary rays of oaks with deciduous 
foliage, appear to have been built up by an aggregation and fusion 
of numerous uniseriate rays. 

5. Traumatic reversions are confined to regions which have been 
severely injured. Occasionally areas where the traumatic effect 
has been slight show an acceleration of the compounding process 
instead of a reversion to ancestral stages. 




fibrovascular bundles. From this phylogenetic relation of the 


inadmissible entirely for the large rays of oak, and that the term 



my sincere thanks to Mr. G. B. 

ifolia. I am also much indebted to Professor 
Jeffrey for suggestions and advice, and to Mr. A. J. Eames 

for material of several wounded oaks. 

Harvard University 
Cambridge, Mass. 



Fig. i. — Quercus nigra: tangential section of the normal adult wood, 
showing the so-called primary and secondary rays; the former are seen to 
consist of large masses of homogeneous ray parenchyma ; X 1 20. 

Fig. 2.— Quercus nigra:* tangential section of the traumatic wood, showing 
compounding mass of small rays; X 120. 

Fig. 3.— Quercus virginiana: tangential section of the normal adult wood, 
showing strong evidences of a compounding process; inclusions of fibers and 
wood parenchyma cells occur conspicuously; X 120. 

Fig. 4.— Quercus virginiana: tangential section of the traumatic wood, 
showing uniseriate non-aggregated rays in the immediate vicinity of a severe 
w <>und; X120. 

Fig. 5 — Quercus densiflora: tangential section of the normal adult wood, 
showing aggregating and fusing mass of small rays; X60. 

3 8o 



— Quercus densifl 


tangential section of the traumatic wood, 



Fig. 7. — Quercus alba: tangential section of the normal adult wood, 
showing highly organized broad homogeneous ray; X 120. 

Fig. 8. — Quercus alba: tangential section of the traumatic wood, showing 
the aggregation and fusion of triseriate and biseriate rays; these rays have been 
produced by the enlargement or growth in diameter of uniseriate rays; X 120. 

Fig. 9. — Quercus alba: transverse section of the normal adult wood, show- 
ing the highly organized type of ray; X120. 

Fig. 10. — Quercus alba: transverse section of the traumatic wood, show- 
ing aggregation of uniseriate, biseriate, and triseriate rays; X120. 

Fig. 11. — Alnus sp.: transverse section of very slightly injured wood; 
compounding rays are absent from the lower half of the section; starting 
abruptly from a line of slightly injured tissue which crosses the middle of the 
section, numerous rays extend outward to the exterior of the stem; X4°- 

Fig. 12. — Quercus densiflora: transverse section of very slightly injured 
wood; the lower half bf the section possesses only uniseriate rays, but start- 
ing from the middle of the section a compounding ray has originated from a 
slight injury to the wood; X60. 













fc ■* 










(with eight figures) 

In the fall of 1909 a species of Achlya that was found to be new 
appeared in cultures from pools around Chapel Hill, N.C. It was sepa- 
rated from other forms and has now been kept under observation in 
pure cultures for nearly a year. It adds one more to the Racemosa 
group proposed by me in 1908. 1 The species may be described as fol- 
lows : 


Achlya caroliniana, sp. nov. — Hyphae rather stout, about 48 m at the base 
and 20 /* near the tip, in strong cultures reaching a length of 1 . 5 cm. Zoospo- 
rangia irregularly cylindric, about 20-30 /* in diameter, often discharging by 
several openings. Oogonia abundant, spherical, smooth, andunpitted; termi- 
nating short, slender branches, 


are racemosely borne on 

the strong main hyphae. Oogonial 
branches generally simple, but often 
giving off one or two branches near 
the base which also terminate in 
oogonia, and, as a rule, are curved 
downward. Oospores generally 1 
or 2, often 3, and very rarely 4 or 
6 (8 were seen twice). They are 
centric, with a diameter varying 
from 18.5 to 26 a*, averaging about 
22 m. Antheridia absent, but hy- 
Pogynous antheridial tubes often 
appear through the basal partition 
exactly as in Achlya hypogyna 
Coker . 

Typical zoosporangia are 
shown in figs. 1-3. In cultures 
that have become somewhat 
foul, the spores may be fully 



'vui, me spores may be fully ' f 

formed, but not discharged. Figs. 1-3 .-Sporangia of various forms; 

In several such cases the spores a11 X33S 


A new species of Achlya. Bot. Gazette 45: 194, 195. 1908. 

[Botanical Gazette, vol. 5° 




were seen sprouting and sending their tubes through the sporangium 
wall, as is normally the case in Alpanes. 

Figs. 4-7 represent the normal arrangement of the oogonia; the 
hypogynal tube will be noticed in fig. 4. In countings from my cultures, 


Figs. 4-7. — Fig. 4, simple oogonial branch terminating in an oogonium; fig. 5' 
oogonial filament with one branch; fig. 6, same with two branches; fig. 7, a branched 
oogonial filament with terminal oogonium containing four oospores; all X335- 

this tube was present only in about one-sixth of the oogonia, but it 
appears in all cultures and is one of the most distinctive characters of the 
species. The appearance of an antheridial tube, not from an antherid- 

ium, but from the vegetative coenocyte, is a most 
singular occurrence, and would seem to be a de- 
generate condition induced by the suppression of 
fertilization. I could find no evidence of fertiliza- 
tion even in cases where a tube was found. In n0t 
a single case was an antheridium seen below the 

The arrangement shown in fig. 6, suggesting the 
three balls of a pawnbroker's shop, occurs so often 
and is so striking as to be one of the best diagnostic 
Fig 8 — E ' cnaracters of the species. The lower branches are 
al oogonium witnTwo" ^^ recurved ( fi g s - 5, 6), but not always (fig. 7>- 
projections on surface; Out of many thousands of oogonia seen, a very 
X335- few had one or two short rounded outgrowths from 


the surface (fig. 8) like those that are characteristic of A. hypogyna. 
In only two cases were intercalary oogonia seen. 

In old cultures the protoplasm becomes condensed and segregated 
into certain restricted areas of the hyphae to form resting fragments, 
which, though not of definite shape, may be regarded as chlamydospores. 

In the Racemosa group, as mentioned above, there may now be 
included A. racemosa, A. racemosa var. stelligera, A. hypogyna, and A. 
caroliniana. The group may be defined as follows: 



few, generally one or two. Antheridia absent, or of suboogonial origin. 
W. C. Coker, Chapel Hill, N.C. 



Until a year ago (June 1909) the saprophytic fungi of the Rocky 
Mountain region included within Wyoming had remained practically 
untouched. It was with great interest, therefore, that the writer began 
the task of making a collection of these fungi under the kindly sugges- 
tion and help of Professor Aven Nelson. The particular region 
studied includes the whole of the Medicine Bow National Forest, together 
with the vast extent of the Laramie Plains. The timber in the forest is 
chiefly lodgepole pine, Englemann spruce, Douglas fir, and balsam, with 
dense growths of aspen on the boundaries. In many places the timber 
is very dense, in consequence of which the humus formed of the needles 
is very thick; the soil in the open timber also is very rich. The whole 
region is well watered by the melting snows and by the numerous 
mountain streams and creeks, resulting in very favorable conditions for 
the growth of the fleshy fungi. The great difference in the altitude, 
which ranges from 2130 to 3900 meters, further aids in the formation of 
many varying conditions, thus giving not only richness in specimens 
but a wealth of species as well. 

In the very precursory examination of the region, the writer was 
astonished at the great quantity and variety of forms. The further 
collections, which it is expected will be made, will no doubt extend the 
1J st greatly. This list, as finally worked out, will be published as a 
w hole, but for the present the following apparently new species only 
a re presented. 

Catathelasma, gen. nov .— Pileus somewhat fleshy, convex, then 
expanded: lamellae very decurrent, somewhat unequal, with acute edges: 


stipe furnished with a ring and of the same substance as the pileus: 
volva large, white, with a thick margin: spores white. 

The ring and volva, together with the very decided decurrent gills (upon 
which character the generic name is based), are telling characteristics of this 
genus of the Leucosporae. 

C. evanescens, sp. nov. — Pileus 13 cm. broad, white, deep cream in 
center, broadly convex to nearly plane, smooth, damp, broadly elliptical 
in outline; margin entire: flesh whitish, compact, thick in center, 
thinner near margin: lamellae very decurrent, short ones intermixed 
with long ones, white, 2-3 cm. wide near margin of pileus, becoming 
narrowed near and on stipe, subdistant, edges acute: stipe very short, 
thick, 1 cm. long, 4 cm. thick, fleshy, hollow, smooth, white; annulus 
delicate, evanescent, situated on stipe just below gills: volva large, 
white, smooth, opening around top leaving a thick even white margin, 
persistent and closely embracing base of stipe : base large, white, bul- 
bous: spores white, smooth, elliptic to fusiform, 14-17- 5^3*5 /*• 
Whole plant when dried becomes a rich ochre with reddish tinge. 

Habitat: Open balsam and spruce woods, occurring singly in sod on thick 
humus; Brooklyn Lake, Wyoming, Snowy Range, alt. 3500 meters, Septem- 
ber 8, 1909, no. 88. 

Clitocybe pruinosa, sp. nov. — Pileus 3.5 cm. wide, piano-con vex- 
to slightly depressed, rich reddish brown over salmon, paler at margin, 
dry, smooth, shining: margin turned down and entire: flesh compact, 
white, tinged with color of cap: lamellae thin, narrowing behind, salmon 
yellow, close, very decurrent, becoming somewhat powdered with 
numerous white spores: stipe fleshy, 5 cm. long, 1 cm. wide, concolorous 
but lighter than cap, hollow, smooth: spores white, spiny, globose, 

Habitat: Pine humus, open pine woods; Foxpark, Wyoming, August 
14, 1909, no. 83. 

Collybia maculata moschata, var. nov.— Pileus fleshy, firm, 6 cm. 
wide, convex to nearly plane, white, glabrous, shining, becoming tinged 
or stained with pinkish red blotches, disk sometimes broken up into 
large polygonal plates; margin turned down and even; flesh white, com- 
pact, tinged with pink just beneath the surface: lamellae whitish, in one 
specimen a faint pink tinge, adnexed to nearly free: stipe stout, firm. 
3 cm. long, 2 cm. wide, swollen in middle, stuffed, becoming hollow, 
curved, narrowed at base, striate, slightly roughened by broken fibers 
spores white, smooth, sometimes with a slight point at one end, 7X4 P- 
It has a strong, almost overpowering odor of musk. 


Habitat: On side of dead lodgepole pine log, clustered; Foxpark, Wyo- 
ming, alt. 2900 meters, August 13, 1909, no. 79. 

Entoloma viridans, sp. nov. — Pileus 3 . 5-5 . 5 cm. broad, fleshy, 
broadly convex, hygrophanous when moist, gray, margin tinged with 
rose pink and disk becoming dull green, or the coloring may be reversed, 
the disk rose pink and margin a dull green, when dry the whole plant 
becomes silky shining: flesh white, becoming dull when dry: margin 
turned down, entire, smooth: lamellae all even, light pinkish yellow, 
becoming a salmon pink color, 2 mm. broad, slightly sinuate, adnate 
then separating, interspaces venose: stipe fleshy, white, pruinate, 
hollow, round, quite bulbous at base, attenuating upward, 4.5 cm. 
l° n g> 1 • 5 cm. wide: spores coarsely war ted, pink, 7X 10 p. 

Habitat: Damp humus; Brooklyn Lake, Wyoming, alt. 3500 meters, 
bank of Nash's Fork, September 3, 1909, no. 119. 

Gloeophyllum ferrugineum, sp, nov. — Pileus hard, corky to woody, 
oblong-dimidiate to flabellif orm, 1-2 X 2-1 2 X 1-1 .5 cm. ; surface 
azonate, strigose-tomentose, scrupose, dark ferruginous to umbrinous; 
margin rather thick, sterile, tomentose, bright ferruginous: context 
corky, homogeneous, bright ferruginous, indistinctly zoned, about 
7 nim. thick; tubes lamelloid, quite decurrent, tomentose, ferruginous 
but paler than margin of pileus, light grayish within, 1-2 mm. broad, 
2-5 mm. deep; edges somewhat thin, tomentose, undulate; tubes 
lamelloid from the first: spores globose-ellipsoid, smooth, hyaline, 

Habitat: On dead lodgepole pine and aspen; Cooper Creek, Wyoming, 
alt. 2800 meters, June 22, 1909, no. 11. 

Clavaria truncata, sp. nov. — Pileate tops bright red, shading into 
reddish orange at top of stipe to dull flesh color at its base : ends truncate, 
convex to plane to somewhat concave, 0.5-3 cm - broad, smooth: whole 
plant to within a few centimeters of base of stipe covered with a white 
bloom, persisting in dried specimens : flesh creamy, spongy : stipe longi- 
tudinally grooved to base, 3-10 cm. long: spores white, 14X7 /*• 

Habitat: Humus soil under balsam and spruce trees; gregarious and 
cespitose, 4-6 in a group; Foxpark, alt. 2900 meters, August 8, 1909, no. 66. 

A plant similar to this is described by Fries as Crakrellus pistillaris, and 
by others as possibly a variety of Clavaria pistillaris, but in a collection of 
tw enty specimens found in entirely different localities not one out of the num- 
ber was found to have either the color or the form of typical Clavaria pistil- 

arrison Lovejoy, University of Wy 



The root-fungi of orchids 

Burgeff 1 has brought together the extended results of his own research 
and those of other students upon the root-fungi of orchids. The volume con- 
tains a comprehensive citation of literature, several tables summarizing results 
of experiments, and discussions of the theory of the mycorhiza question. By 
way of introduction, Burgeff defines his use of the word symbiosis as that 
relation of two symbionts in which one aids the other in any way, if only to 
make existence possible under conditions otherwise impossible. After union, 
the two organisms form a new organism, a unit, which takes up the struggle 
for existence under new conditions, each being a body member. Such is claimed 
to be the condition of the orchids and their root-fungi. The two sections of 
the work are as follows: 

i. The study of the fungus independent of the plant— On a culture medium 
of agar and rain water with a slight trace of starch, 29 root-fungi, aseptically 
obtained from native and tropical orchids, were grown (holosaprophytic ones 
were unsuccessful). Such species of Orcheomyces, as the author chooses to call 
the fungi for convenience, are described in detail as to their structure and 
behavior in the culture. For the first time a study of their enzymes has been 
made, and the endophytes were grouped accordingly on the basis of their 
biological relations to the orchids. In general they have thin-walled, regularly 
septate mycelia; the hyphae are sharply differentiated into Langhyphen that 
branch little and show unlimited growth in one direction, and Kurzhyphen that 
are of smaller caliber and arise at regular intervals and whose cells under 
certain conditions are transformed into spores (Sporentrager) or absorptive 
hyphae (Sanghyphen). In old age spores and hyphae contain a fatty oil. 
Chains of hyaline or slightly colored spores may unite into loose clusters or 
closer sclerotia-like groups. All fungi fuse or anastomose in some manner. 
The hyphal cells contain two to ten nuclei, but the spores contain only two 
nuclei. Spiral knots initiated in cultures by the surface tension of a drop oi 
water at the tip of the hypha are comparable to those in the root cells developed 
in response to the pressure of the resisting plasma membrane to the penetration 
of the hypha. No sexual reproduction was observed. 

* Burgeff, Hans, Die Wurzelpilze der Orchideen; ihre Kultur und ihr Leben in 
der Pflanze. 8vo. pp. 207. pis. 3. figs. 38. Jena: Gustav Fischer. 1909. 

, Zur Biologie der Orchideen Mycorhiza. pp. 66. Inaug. Diss. Jena. 

Gustav Fischer. 1909. 




Of the carbohydrate culture media, starch, maltose, and saccharose were 
best; glucose and dextrin next; and glycerin poorest. Correspondingly, all 
fungi possessed diastase and emulsin; some invertase and maltase; some 
aesculin; one tyrosinase; and only one cytase. In humus decoctions all 
grew well. 

The endophytes cannot assimilate free nitrogen. They belong to Ben- 
necke's category of "Ammon-nitrit-nitratpilze," and grow luxuriantly, form- 
ing spores on ammonium salts, ammonium nitrate holding first place. Organic 
N compounds were varyingly well assimilated. Peptone produced splendid 
growth, and salep furnished sufficient N for all to grow. All fungi possess 
proteolytic enzymes. 

The production of acid in assimilation is slight, and in general, in media 
containing asparagin, peptone, or urea, is in proportion to the intensity of the 
growth. Orchid fungi require to a high degree atmospheric oxygen, dying 
in long-continued anaerobic cultures. The formation of spores and spore 
scferotia depends upon the concentration of the medium, its exhaustion, and 
the amount of assimilation products present. Salep always stimulated spore- 
formation, as did increased transpiration. 

2. The study of the plant and the fungus. — For the biological relations of 
the plant and the fungus, Burgeff chooses the terrestrial orchid Epidendrum 
(dickromum?) and the hybrid epiphyte Laelio-Cattleya. Without a fungus, 
seeds of Epidendrum, on a culture medium of rain water, 2 per cent salep, and 
1 -5 per cent agar, did not germinate. With ten different fungi they germinated 
within 25 days. The embryo did not become green until infected. Thinned 


cultures. The stages 

Cypripedium type of development described by Bernard. Infection takes 
place through the suspensor at first, and later through the root hairs, passing 
by means of the Durchlasszellen into the cortex. The hyphae of the sub- 
epidermal layer (Pilzwirtzellschicht) are never digested, but Eiweisshyphen 
appear in the digestive layer beneath. The cytological facts agree with the 
phenomena of digestion, where the remains (a clump) are surrounded by a 
cellulose layer (Haut), and the nucleus assumes the resting stage, ready to 
^gest new hyphae. Clumps may become several-layered. 

Burgeff found that the seeds of Laelio-Cattleya, wettable only after three 
or ( our days, could germinate, become green, and attain considerable differ- 
entiation (stomata at apex, rhizoids at base, leaf primordia) in three or four 
months. A resting period of a year is necessary before growth continues, for 
which also a fungus is required. Just before infection, the oil in reserve in 
the cells n ear the suspensor is transformed to starch, which disappears, however, 
^ soon as the fungus penetrates the cells. Both embryo and fungus show 
evidence of growth, the latter by the formation of Eiweisshyphen (homolo- 

spores and hyphal 

germination in Laelio-Cattley 



Burgeff obtained its development, in 0,33 per cent cane sugar with mineral 
salts on agar, in the dark, to the differentiation of papillae, and it lived ten 
months. Further development either in light or dark required the fungus. 
While in the light the orchid deve loped normally, in the dark it elongated and 
resembled the Epidendrum which grew in the weaker concentration of salts. 
Burgeff concludes that physio logi cally the behavior of the fungus is alike 
in the germination of the epiphyte and the terrestrial orchid, and that the 
appearance of chlorophyll in the epiphyte is an adaptation to its life in the 

Seedlings with the fungus in absence of C0 2 in the dark grew, but developed 
no root, while in the light the growth was normal. Experiments on the physi- 
ology of nutrition, where fungus and seedling were grown together, showed the 
best N sources to be ammonium chlorid and nitrate. Although asparagin was 
favorable for the fungus in free culture, the plant could not grow in it. In 
N-free media there w r as no development. Lactic acid (2 per cent) in minera 
salt agar culture caused both plant and fungus to grow well, Laelio-Cattleya 
was found to harbor some fungi (17 tested) unable to stimulate its germination- 

Burgeff devotes 50 pages to the histological processes in the growing 
plants. These detail the places of entrance, position of the fungi, peculiar 
features in the plant cell (spores and clumps, Klumpenbildung), the emission 
of hyphae to the substratum, etc. Most of the fungi have mycelial connections 
with the outside substratum, therefore anatomically there is nothing to prevent 
ascribing to such hyphae the function of conduction of soluble materials. The 
author explains the unwettability of the seeds as due to the Luftblaschen in the 
netted testa itself, and to the air between the testa and embryo (comparable 
to the condition in the lycopod spores). Such unwettability hinders the pas- 
sage of spores through the soil, contrary to the theory of Koch and Lustner, 
but is of advantage in preventing the clinging together of seeds in the capsule, 
therefore an adaptation to their dissemination by wind. Also because the seeds 




xnaubtion 01 imb suusumv^. 

in the probable steps of development of 
the symbiosis by way of parasitism, but would speak of the association not as 
a "maladie bienfaisante," but as of "einem gliicklichen Zusammentreffen 
verschiedener Umstande." He bases this remark on the following facts: the 
fungus is harmless, its enzymatic qualities separating it froi 


parasites with toxic qualities; 

ted fungu 

penetration it nevertheless allows; its fitness for infection is seen in the 
Durchlasszellen of the embryo. A mutualistic symbiosis demanded by the 



to obtain from the soil which enable it to form spores. As to the materials 
of exchange between the components, the question is left unanswered; but the 
idea of conduction of mineral salts is favored, because of the results 01 the 


cultural experiments and because of the detailed morphological and anatomical 
features of both symbionts. The fungus causes the conversion of starch into 
sugar by its diastase. Its function results from its enzymatic quality, which, 
with the solution of carbohydrates in the plant cell, induces the development 
of the seed, not by bringing soluble materials to the cell, but by transforming 
substances already there. Burgeff suggests here the unproven fact of diffu- 
sion of the diastatic enzyme out of the fungal hypha through the Plasmahaid 
into the plant cell. This may also occur in the substratum from the emission 
hyphae. The osmotic relations arising from the sugar solutions could account 
for the absorption of water, but if nutritive salts are absorbed from the fungi 
from outside, a rapid change in permeability and adjustment of pressures at 
just the proper time to seize the salts brought by the fungus must take place. 
On the whole, the relations between the plant and the mineral salts of the 
soil are of striking importance for the origin and maintenance of the orchid 
symbiosis. Although the structures show a gain in nitrogenous substances, 
the habitats of orchids, and cultural experiments exclude the possibility of free 
N-absorption. No anatomical features can prove the absorption of organic 
carbohydrates; although diastase and emulsin are common to all fungi, material 
for the action of the former is lacking in the soil, and we are in ignorance con- 
cerning the substance in the soil digested by the latter. Any substance taken 
up by the plant, either through its roots or by means of the fungus, must first 
be made soluble by the fungus itself, or by its exoenzymes in the substratum. 

Grace L. Clapp. 


Farm weeds. — The preparation of a scientific manual for the use of the 
ordinary layman is admittedly a difficult task, but it has been successfully 
accomplished by Clark and Fletcher, 2 whose volume upon farm weeds is the 
best that has yet appeared upon this subject. The remarkable simplicity 
without the sacrifice of scientific accurracy is due largely to the splendid 
ability of the late Dr. James Fletcher, who thus adds one of the latest of his 
many valuable contributions to botany and agriculture. More than 200 of 
the more troublesome weeds of Canada are arranged according to modern 
botanical classification, with very complete scientific and common synonyms, 
and briefly but accurately described in non-technical language. Special 
attention is directed to the characteristics which make the various plants 
troublesome as weeds, and careful directions are given for the most prac- 
ticable and successful methods of control and extermination. 

The most valuable aid to the recognition of different species is a series 
°f 76 full-page plates, colored with the greatest accuracy. They include 


edition. 8vo. pp. lg2 . ph. 7 6. Ottawa: Department of Agriculture, Dominion of 
Canada. 1909. $ Ii0 o (single copies only, for sale by Superintendent of Stationery, 
Government Printing Bureau, Ottawa). 


representatives, in natural size and also much enlarged, of the seeds of ioo 
of the most troublesome species. These illustrations are from the water- 
color sketches of Norman Criddle, and will enable the farmer to identify- 
readily most of his plant enemies, either while they are growing in the field 
or while polluting his seed grain. 

The book is well printed and strongly bound, which, together with its 
other admirable qualities, makes it a valuable addition to the literature of 
economic botany. — Geo. D. Fuller. 

A text-book of pharmacognosy. — A fourth edition of Kraemer's Text- 
book has appeared.^ It is "intended for the use of students of pharmacy, as a 
reference book for pharmacists, and as a handbook for food and drug analysts." 
Such a statement indicates that the volume does not fall within the province 
of a botanist for review, and yet the material presented is of great interest to 
botanists. Part I (pp. 222) is entitled "Botany," and comprises a presenta- 
tion of all the great groups, "outer morphology of angiosperms," "inner mor- 
phology of the higher plants," "classification of angiosperms yielding vegetable 
drugs," and "cultivation of medicinal plants." Of course this is botany for 
the pharmacist, and Dr. Kraemer is in a position to know what the pharma- 
cist needs. Only the first chapter, dealing with the great groups, really per- 
tains to the non-pharmaceutical botanist. Perhaps it makes no difference 
to the students concerned, but the very antique flavor of the presentation of 
the great groups is somewhat surprising to the modern morphologist. Part 
II (PP- 383) is entitled "Pharmacognosy," and deals first with crude drugs, 
and then with powdered drugs and foods. Part III (pp. 88) is entitled 
"Reagents and technique"; and part IV (pp. 38) deals with "Micro-analysis." 

The volume is certainly a thesaurus of information for the pharmacist, 
and doubtless will have great influence upon the progress of pharmacognosy 
in this country. There is abundant evidence, also, of an immense amount 
of painstaking labor on the part of the author, who is to be commended for 
his many years of faithful effort to organize and advance his subject. — J, M. C. 

A naturalist in the Bahamas. — Under this title a memorial volume in honor 
of Dr. John I. Northrop has appeared.* The botanical papers are as fol- 
lows: "Flora of New Providence and Andros (Bahama Islands)," by Alice R 




(reprinted from Bull. Torr. Bot. Club 17:1890); "Notes on the plant distri- 

by John 


pp. viii+888. 

Philadelphia and London : J. B. Lippincott Company. 1910. $5.00. 

4 A naturalist in the Bahamas. John I. Northrop. Oct. 12, 1861 — June 25, 



field Osborn. pp.281. New York : The Columbia University Press. 1910. S2.50 


"A study of the histology of the stem of the wax plant, Hoya carnosa (L.) 
R. Br.," by John I. Northrop.— J. M. C. 


Prothallia of Lycopodium. — Bruchmann 5 has completed the life histories 
of L. clavatum, L. annotinum, and L. Selago, with an account of the growth 
of the prothallia from the germination of the spores to the maturing of the 
sex organs, and with the study of the embryo development of L. Selago. Pre- 
viously the germination of only three species was known, all with a pro thallium 
containing chlorophyll and belonging to the cernuum type: L. salakense, 
independent in development and free from a fungus; L. cernuum; and L. 
inundatum, dependent for complete development upon an endophytic fungus. 

The germination of the spores was secured by sowing them in the forest 
where they were found. This was done in two ways: (1) by putting spores 
and species of fresh strobili into flower pots in potting soil (peat, leaf mold, 
manure, sand) and forest soil and sinking them in the ground; and (2) by 
mixing spores with forest soil and burying them in holes 10 cm. deep. The 
germination of L. Selago occurred in 3-5 years after sowing; that of L. clavatum 
and L. annotinum in 6-7 years. Prothallia bearing mature sex organs appeared 
first in L. Selago 6-8 years after sowing; in L. clavatum and L. annotinum in 
12-15 years. Practically all the spores of a sporangium of L. Selago germi- 
nated, but only about 5 per cent of those of other species. All three species 
develop independently a five-celled chlorophyllose pro thallium, after which 
a fungus is necessary to complete the development. L. clavatum and L. 
annotinum develop in the same way, while L. Selago forms a type by itself. 

The first division, an irregular one, cuts off a small lens-shaped cell at the 
base of the spore. Although containing food substances, it is poor in cyto- 
plasm and never enlarges. Bruchmann considers it a rudimentary rhizoidal 
cell, homologous with the first rhizoid of the true ferns. The second division, 
occurring after the spore coat bursts, extends an oblique wall from base to 
apex, bent toward the rhizoidal cell. While in the clavatum type this wall 
divides the prothallium into hemispheres, in the Selago type it is sharply bent 
away from the rhizoidal cell, making the resultant cells unequal. The differ- 
ence in direction determines the type of prothallium. The basal cell next to> 
the rhizoidal cell never divides. Its sister cell cuts off an inner wedge-shaped 
cell and an outer apical cell. A fourth wall divides the wedge into an outer 
Peripheral and an inner or central cell. In L. Selago the wedge-shaped cell 
is considered the first segment cut off from the apical cell. 

From this point Bruchmann divides the growth of the prothallia into 
three periods, depending upon change in direction and manner of growth. 

5 Bruchmaxn, H., Die Keimung der Sporen und die Entwicklung der Prothal- 
tien von Lycopodium clavatum, L. annotinum, und L. Selago. Flora 101:220-267. 

fits. 35- 



L. Selago resembles closely L. Phlegmaria in development, while L. clavatum 
and L. annotinum are alike. The entrance of the fungus at the peripheral 
or basal cell stimulates the apical cell to division. It cuts off from its two sides 
alternately five or seven segments in the L. clavatum type, and fewer in L. 
Selago. This closes the first period. 

Extensive growth of both the prothallium and the fungus characterizes 
the second stage. Rapid radial growth at the apex results in a pear-shaped 
prothallium. The fungus in the two lycopod types of prothallium represents 
two species. In L. clavatum it forms a peripheral digestive mantle surrounding 
the central mass of storage cells. Limited from the central cells by a palisade 
layer, it becomes entirely intercellular. The fungus of L. Selago resembles 
in habit that of L. Phlegmaria and fills the entire prothallium posterior to 
the apical region except the epidermis. The sister cell of a rhizoid, infected 
by a branch of the mycelium from the hypodermis, serves as an " Expeditions- 


zelle." From it the hypha passes out to the substratum, spraying out into 
fine branches. To this cell Bruchmann attributes the function of chemically 




meristematic cells in L. clavatum surround the axial conductive cells, coexten- 
sive with the former storage cells. Reproductive organs develop on the margin, 
antheridia preceding archegonia. Such prothallia may live twenty years. 
During this period the fungus forms spores. In L. Selago the attempt to gain 
dorsiventrality causes extensive elongation of the prothallium, which at the 
soil surface develops chlorophyll. The apical growth becomes marginal, the 
central tissue acting as a storage region. The outer layers on one side develop 
paraphyses and sex organs; on the other, vegetative cells containing the 
fungus, which always remains intracellular. 

The embryo-development of L. Selago agrees in detail with that of L. 
Phlegmaria. The root appears last, later than in L. clavatum and L. annotinum. 
The foot is smaller than that of L. clavatum and of L. annotinum, but less 
papillate than that of L. Phlegmaria. Its continued growth upward bursts 




Variation in timothy. — An important contribution to the subject of 
secular variation has been made by Clark, 6 who has studied the variation 
in height, weight of forage produced, earliness of bloom, and duration of the 
period of bloom in timothy (Phleum pratense). Data were secured during 
three successive years on 3505 plants representing 163 pedigrees derived from 
2 2 different states of the United States. As there can be no doubt that timothy, 
like many other plants which have been studied, consists of a number of distinct 
hereditary forms or bio types, the extent of variation, which was found to be 

6 Clark, C. F., Variation and correlation in timothy. Bull. 279, Cornell Uni- 
versity, pp. 301-349. figs. 111-150. July 19 10. 


very great in all characters studied, may not be considered as having great 
significance, since the bringing in of still other pedigrees from other sections 
would doubtless have increased the range of variability. No perceptible cor- 
relation was found between earliness and height of the plants or between 
duration of bloom and height of plants. There appeared to be a slight nega- 
tive correlation between the duration of bloom and weight, but this was very 
slight and possibly not significant. Between weight and height, as might be 
expected, there was considerable positive correlation, ranging from 0.274=*= 

0.011 to 0.718=1=0.006. By securing data covering three years from the 
same series of plants, an interesting new relation has been developed, namely, 
the correlation between the condition of plants in one year as compared with 
the same plants in succeeding years, and for this correlation the author gives 
the name " coefficient of place-variation." This measures the extent to which 
an individual, found to have a given rank with respect to a variable character 
in one year, may be expected to hold the same rank in succeeding years, and 
is a very important consideration from the standpoint of the practical breeder. 
The correlation coefficients found ranged from 0.382=^=0.010 to 0.585=1=0-008. 
The lowest correlation was found in comparisons between non-consecutive 
years, as when 1905 was compared with 1907. This would naturally be 
expected, since there are more disturbing factors in two years than in one. 
These coefficients are considered rather low, and are taken to indicate the 
importance of comparing individuals during several years as a safe basis for 
selection in economic breeding, since there are very good chances that an 
individual observed to be superior in one season may be inferior in succeeding 
seasons.— Geo. H. Shull. 

Seeds of horseradish.— It is a well known fact that the horseradish (Cock- 
learia Armoracia) is generally sterile, though it produces a great abundance 
of flowers and not infrequently produces capsules. Brzezinski 7 has induced 
the development of seeds by removing a circle of the bark from the upper 
Portion of the root a short distance below the collum. Plants so treated 
produced a considerable number of good seeds, and in one year (1908) he secured 
x 5oo seeds. Of 50 seeds sown in 1907, 30 produced plants, most of which 
succumbed to disease, but 9 of which grew to maturity. From the same (1906) 
crop, 200 seeds planted in 1908 produced only 20 seedlings, thus apparently 
indicating the rapid loss of vitality of the seeds. Only 6 of these reached 
maturity. These 15 mature seedling plants of the horseradish were not uni- 
form, but were referable to two types, neither of which agreed with the char- 
acters of the parent. The ordinary horseradish is intermediate between these 




considerable number of seeds, even without the operation which induced seed- 

7 Brzezinski, J., Les graines du raifort et les r6sultats de leurs semis. Bull. 
Ac *d. Sci. Cracovie, Session of July 5, 1909. pp. 392-408. pis. 12-15. 

394 BOTANICAL GAZETTE [november 

production in the parent. Two hypotheses are offered to account for the 
appearance of these two types among the seedlings: (i) that they are muta- 
tions produced as a result of traumatism in accordance with the views of 
Blaringhem; and (2) that the ordinary horseradish is not a natural species 
as generally believed by taxonomists, but a hybrid, and that the two types 
of offspring produced from the seeds are partial or complete returns to its 
parent types. The author inclines to the latter view, and would interpret 
the sterility of the horseradish as due, not to the accentuated development 
of fleshy roots, but to a weakening of the sexual development, not infrequently 
found in hybrids. The reviewer is inclined also to the latter interpretation, 
and would point out the important bearing the author's method of securing 
seeds of the horseradish may have in its application to other sterile hybrids. 
Many experiments have been terminated by the failure of hybrids to produce 
seeds. It may be that some of these cases will yield to methods of treatment 
similar to that employed by Brzezinski in securing seeds of horseradish. 

Geo. H. Shull. 

The ecology of conifers. — Stopes and Moss have discussed the xerophyt- 
ism of conifers, and now Groom 8 considers a number of their ecological 
features. In the introductory statement three problems are outlined: the 
cause of their xerophytic foliage and tracheidal wood, the cause of their 
survival in competition with dicotylous trees, and the cause of the suppression 
of many forms in past ages. Groom correctly concludes that not all conifers 
are xerophytic, in spite of their xerophytic leaf structure, calling attention to 
von Hohnel's demonstration of high transpiration in the larch, and to his 
own experiments which show that coniferous wood, in spite of its tracheidal 
structure, may conduct water with a rapidity equal to that of a rapidly tran- 
spiring dicotylous tree. Attention is called to the fact that the aggregate 
leaf surface of a coniferous tree may exceed that of a dicotylous tree, because 
of the immense number of leaves. Indeed, Groom regards the xerophytic 
structure of the leaf as a necessity in view of the great amount of exposed 
surface, and he applies the term "architectural xerophytism" to xerophytism 
that is dependent upon the organization of the plant rather than upon the 
direct influence of external factors upon the organs in question. In opposi- 
tion to Stopes, Groom regards the tracheidal nature of the wood as a feature 
of advantage rather than a feature necessitated by heredity, and notes that 
similar wood tends to occur in various evergreen dicotyls. The extinction 
of many conifers of past ages is attributed to their imperfect acclimatization, 
to the fact that they have a great number of insect and fungus enemies, 
and to their relatively slight power to react advantageously to new conditions. 



8 Groom, Percy, Remarks on the oecology of Coniferae. Annals of Botany 
24:241-269. 1910. 


to those that are sufficiently humid to permit the development of luxuriant 
mesophytic forests. Groom's paper is most suggestive, and adds considerably 
to our knowledge concerning the difficult problem of coniferous xerophytism. 

Henry C. Cowles. 

Nutrition of the embryo in Labiatae. — Billings 9 has investigated the 
nutritive mechanism associated with the embryo sac of certain Labiatae, 
a subject that deserves more attention from morphologists. The ordinary sac 
which is oval or elliptical in longitudinal section, and which encroaches uni- 
formly upon the surrounding tissues, has come to be regarded as the more 
or less fixed " type" of angiospermous sac. Among the Sympetalae especially, 
however, a much more complex nutritive mechanism has begun to be uncovered, 
including special digestive layers and special absorptive regions of the sac, the 
latter usually taking the expression of tubular haustorial extensions. Billings 
investigated 15 species of Labiatae, representing 14 of the most representative 
genera. The results were uniform enough and differed enough from other 
sympetalous groups investigated to indicate that such structures may be of 
taxonomic and even of phylogenetic value. For example, the Scrophulariaceae 
previously described usually have a well developed digestive layer 
("tapetum"), in addition to haustorial extensions of various kinds; but 
the Labiatae lack the special digestive layer. There are three features com- 
mon to the species studied, and possibly to the whole family, to which the 
author calls attention: the micropylar haustorium (more or less extensively 
developed), the much-elongated suspensor, and the antipodal canal or process. 
Salvia is an exception to this statement, for it has a short suspensor and no 
micropylar haustorium; and the two species investigated "are unique in 
having two haustorial outgrowths, one coenocytic and one composed of ordi- 
nary endosperm tissue" (these haustoria are in addition to the well developed 
antipodal canal). The author thinks that such variations from the general 
conditions as are shown by Salvia "suggest a taxonomic rearrangement." 
J. M. C. 

Correlation in oats. 


of head, number of grains per head, and average weight of grains in a variety 

of oats growing at Dickinson, North Dakota. The examination of 1000 
plants discovered decided negative correlations (—0.595=4=0.013, 

o. sn 

°°i5, and —0.404 ±0.01 7) between the weight of grains and number of 
grains per head, weight of grains and length of head, and between weight of 
grains and length of culm. He reaches the conclusion that in selecting the 
heaviest grains in this variety, the breeder selects plants somewhat below the 

9 Billings, F. H., The nutrition of the embryo sac and embryo in certain 
Labiatae. Kansas Univ. Bull. 3:67-83. pis. 11-14- i9°9- 

10 Waldron, L. R., A suggestion regarding heavy and light seed-grain. Amer. 
Nat. 44:48-56. 1910. 

396 " BOTANICAL GAZETTE [November 

average height, with shorter heads and fewer grains, thus emphasizing the 
importance of selecting the superior plants instead of the superior individual 
grains. As this variety of oats was undoubtedly a mixture of several distinct 
biotypes, it does not follow that the same mathematical results would be 
found in other varieties composed of different mixtures. The variety with 
which Waldron worked may have contained a short-headed, short-culmed, 
heavy-grained biotype. In some other mixture the ' heavy-grained biotype 
might have longer culms, longer heads, and more numerous grains, and it 
would then give a positive correlation where Waldron found a negative 
correlation, but this does not lessen the importance of the conclusion reached 
that the individual plant and not the individual grain is the proper unit of 
selection. — Geo. H. Shull. 

Anatomy of the seedling of Trapa. — A short paper by Queva 11 on the 
curious seedling of Trapa natans recalls the case of "caulicle" vs. " radicle," 
the question of the importance of the root as a primarily essential part of a 
seed plant. The author confirms the observations of previous investigators 
concerning the marked inequality of the cotyledons, the negative geotropism 
of the caulicle (which he prefers to call the hypocotyledonary axis), and the 
presence of internal phoem in the stem and leaves. His own investigation 
has resulted in the discovery of this internal phloem in the hypocotyledonary 
axis and in the petiole of the cotyledon. Although he finds in the very tip 
of the hypocotyledonary axis a vascular condition which is peculiarly root- 
like (two xylem points alternating with two small groups of phloem), yet he 
thinks it is not root, because (i) there is no rotation of the strands in succes- 
sive levels from the tip of the organ to the cotyledonary node; (2) the xylem 
points are too near the periphery of the cylinder to look like root poles; and 
(3) the whole organ is covered by epidermis, except at the spot where the 
suspensor was attached. The growth of the caulicle, or hypocotyledonary 
axis, is limited; the roots strike out from its side, their vascular strands being 
inserted on certain metaxylem elements discernible in cross-section on one 
side of the vascular cylinder. — Sister Helen Angela. 

Sporophylls of Selaginella. — Sykes 


and a degree of complexity that have not attracted attention heretofore. 
A few of the more representative species are described and the different 
forms of the sporophyll are pointed out as "special adaptations for the secure 
protection of the sporangia." In many sporophylls there is a well developed 
air cavity in the base, and the authors suggest "that they recall the mucilage 


C, Observations anatomiques sur le Trapa natans L. Compt. Rend. 
Assoc. Fran. Av. Sci. 1909. Congres de Lille, pp. 512-517. Jigs. 2. 1909. 

13 Sykes, M. G., and Stiles, W., The cone of the genus Selaginella. Annals of 
Botany 24:523-536. pi. 41. 1910. 



cavities of Lycopodium and the parichnos of fossil genera." In some cases the 
sporophylls are provided with more or less conspicuous dorsal outgrowths, 
which have been noted heretofore only in a casual way, but not definitely 

described. Four types 

(i) with well developed 

dorsal flap extending freely downward and protecting the young sporangium 
immediately below it (as S. rupestris) ; (2) with no dorsal flap, the sporophyll 
being flat and the sporangium exposed (as S. spinosa) ; (3) with a well developed 
dorsal projection which is not free but decurrent (as S. helvetica) ; and (4) a 
series in which the dorsal outgrowth is gradually reduced and lost, each sporo- 
phyll more and more completely infolding the sporangium below (as S. flabel- 
lata to S. apus) — J. M. C. 

Morphology of Callitris. — Saxton, x 3 in continuing his studies of gym- 
nosperms, has given an account of Callitris, an Australasian genus of about 
a dozen species. The one chiefly studied was C. verrucosa, but the results 
doubtless apply to the genus as a whole, since the species are very closely allied. 
The sporophylls are in alternating whorls of three, each microsporophyll bear- 
ing three sporangia, and each of the upper megasporophylls bearing about 
15 ovules, the six sporophylls of the ovulate strobilus producing about 60 
ovules. The cells of the mature female gametophyte are all binucleate or 
multinucleate. The archegonia occur in a single group of about 20, never at 
the apex of the gametophyte, but along the inner side of the pollen tube near 
its apex. If two pollen tubes are present, two such groups are organized. 
The proembryo completely fills the archegonium, the arrangement of cells 
being variable. More than one embryo is formed from a proembryo, and the 
first two walls are longitudinal, the mature embryo being dicotyledonous. 
The claim is well substantiated that Callitris and Widdringtonia are two distinct 
genera, but that they should constitute a separate tribe (Callitrineae) coordi- 
nate with Cupressineae is not so clear. — J. M. C. 

Structure of Mitrospermum. — Mrs. Arber 1 * has investigated the structure 
of the platyspermous seed described by Williamson in 1877 as Cardiocarpon 
compression, and occurring in the British Lower Coal-measures. The outer 
fleshy (forming the wing-like extension), stony, and inner fleshy layers are 
recognized and described. Two opposite vascular strands arise from the 
expanded bundle beneath the nucellus and traverse the outer fleshy layer in the 
Principal plane of the seed, probably continuing almost the whole length of the 
seed. The nucellus seems to have been entirely free from the integument, and 
in one seed a tissue within the embryo sac was observed, consisting of "irregu- 
lar roundish cells/' which of course represent the female gametophyte. The 

13 Saxton, \Y. T., Contributions to the life history of Callitris. Annals of Botany 
2 4: 55 7-569. pis. 4S, 46. 1910. 

I4 Arber, Agnes, On the structure of the paleozoic seed Mitrospermum com- 
Pressum (Will.). Annals of Botany 24:491-509. pis. 37~39- fiP- 2 - l 9™- 

398 BOTANICAL GAZETTE [november 

conclusion is reached that the seed deserves to be removed from Cardiocarpon, 
chiefly on account of its vascular structure, and therefore a new genus Mitro- 
spermum is proposed. Whether the seed belongs to Cordaites or not was not 
determined, for the platysperm character can no longer be used as an indica- 
tion of that group. Sections of unattached seeds must continue to be made, 
but there is far greater need of sections of attached seeds, for these will probably 
solve the puzzling embryo situation attributed to paleozoic seeds. — J. M. C. 

Phototropism. — Nordhausen 1 * offers more evidence against the lens 
theory of phototropic perception. He finds that leaves of Begonia with killed 
epidermis assume the normal light position, the palisade cells being the per- 
ceptive organs. He says that "the epidermis as w r ell as its papillose character 
are not necessary for light perception. " He finds a great difference between 
the sensitiveness of the two halves of the leaves of Tropaeolum, which renders 
them unsuitable for comparing the effect of light on the wet and dry halves. 
This plant and method have furnished Haberlandt with his best evidences 
for the lens theory. After offering this significant evidence against the theory, 
he states that Haberlandt's reply to his former criticism has not rendered 
that criticism any less applicable. He also holds that the evidence offered 
in Haberlandt's later papers is not of a sufficiently critical nature to give 
the theory any support. — William Crocker. 

Carbon dioxid as a fertilizer. — A Berlin company has placed a product 
on the market known as "Germanol," which consists of an earthy mixture 
containing about 18 per cent calcinated soda. The company attributes the 
virtue of this mixture to an increased porosity of the soil following an increase 
in the proportion of carbon dioxid. Mitscherlich/ 6 however, is of the opinion 
that if such a mixture has any value it must be attributed to the action of the 
carbon dioxid in increasing the solubility of various difficultly soluble soil 
substances. His comprehensive tests show that increasing the carbon dioxid 
content of the soil does not result in an increase of plant product; that there 
is always sufficient carbon dioxid in the soil to render mineral food available; 
that an increase in the carbon dioxid in the soil does increase the solubility 
of difficultly soluble substances, but that such increase is superfluous so far 
as any advantage to the plant is concerned. — Raymond H. Pond. 

Rhizophore of Selaginella. — Worsdell 1 ? has used an investigation of the 
rhizophore of Selaginella as the basis of a discussion of the ultimate morpho- 

** Nordhause