THE CHEMICAL BASIS OF PLANT FORMS.[1]

By HELEN C. DE S. ABBOTT.

The succession of plants from the lower to the higher forms will be

reviewed superficially, and chemical compounds noted where they

appear.

When the germinating spores of the fungi, _myxomycetes_, rupture their

walls and become masses of naked protoplasm, they are known as

plasmodia. The plasmodium _Æthalium septicum_ occurs in moist places,

on heaps of tan or decaying barks. It is a soft, gelatinous mass of

yellowish color, sometimes measuring several inches in length.

The plasmodium[2] has been chemically analyzed, though not in a state

of absolute purity. The table of Reinke and Rodewold gives an idea of

its proximate constitution.

Many of the constituents given are always present in the living cells

of higher plants. It cannot be too emphatically stated that where

"biotic" force is manifested, these colloidal or albuminous compounds

are found.

The simplest form of plant life is an undifferentiated individual, all

of its functions being performed indifferently by all parts of its

protoplasm.

The chemical basis of plasmodium is almost entirely composed of

complex albuminous substances, and correlated with this structureless

body are other compounds derived from them. Aside from the chemical

substances which are always present in living matter, and are

essential properties of protoplasm, we find no other compounds. In the

higher organisms, where these functions are not performed

indifferently, specialization of tissues is accompanied by many other

kinds of bodies.

The algæ are a stage higher in the evolutionary scale than the

undifferentiated noncellular plasmodium. The simple _Alga

protococcus_[3] may be regarded as a simple cell. All higher plants

are masses of cells, varying in form, function, and chemical

composition.

A typical living cell may be described as composed of a cell wall and

contents. The cell wall is a firm, elastic membrane closed on all

sides, and consists mainly of cellulose, water, and inorganic

constituents. The contents consist of a semi-fluid colloidal

substance, lying in contact with the inner surface of the membrane,

and, like it, closed on all sides. This always is composed of

albuminous substances. In the higher plants, at least, a nucleus

occurs embedded in it; a watery liquid holding salts and saccharine

substances in solution fills the space called the vacuole, inclosed by

the protoplasm.

These simple plants may be seen as actively moving cells or as

non-motile cells. The former consist of a minute mass of protoplasm,

granular and mostly colored green, but clear and colorless at the more

pointed end, and where it is prolonged into two delicate filaments

called cilia. After moving actively for a time they come to rest,

acquire a spherical form, and invest themselves with a firm membrane

of cellulose. This firm, outer membrane of the _Protococcus_

accompanies a higher differentiation of tissue and localization of

function than is found in the plasmodium.

_Hæatococcus_ and plasmodium come under the classes algæ and fungi of

the Thallothyta group. The division[4] of this group into two classes

is based upon the presence of chlorophyl in algæ and its absence in

fungi. Gelatinous starch is found in the algæ; the fungi contain a

starchy substance called glycogen, which also occurs in the liver and

muscles of animals. Structureless bodies, as _æthalium_, contain no

true sugar. Stratified starch[5] first appears in the phanerogams.

Alkaloids have been found in fungi, and owe their presence doubtless

to the richness of these plants in nitrogenous bodies.

In addition to the green coloring matter in algæ are found other

coloring matters.[6] The nature[7] of these coloring matters is

usually the same through whole families, which also resemble each

other in their modes of reproduction.

In form, the algæ differ greatly from filaments or masses of cells;

they live in the water and cover damp surfaces of rocks and wood. In

these they are remarkable for their ramifications and colors and grow

to a gigantic size.

The physiological functions of algæ and fungi depend upon their

chemical differences.

These facts have been offered, simple as they are, as striking

examples of chemical and structural opposition.

The fungi include very simple organisms, as well as others of

tolerably high development, of most varied form, from the simple

bacillus and yeast to the truffle, lichens, and mushrooms.

The cell membrane of this class contains no pure cellulose, but a

modification called fungus cellulose. The membrane also contains an

amyloid substance, amylomycin.[8] Many of the chemical constituents

found in the entire class are given in _Die Pflanzenstoffe_.[9]

Under the _Schizomycetes_ to which the _Micrococcus_ and

_Bacterium_[10] belong are found minute organisms differing much in

form and in the coloring[11] matters they produce, as that causing the

red color of mouldy bread.

The class of lichens[12] contains a number of different coloring

substances, whose chemical composition has been examined. These

substances are found separately in individuals differing in form. In

the _Polyporus_[13] an acid has been found peculiar to it, as in many

plants special compounds are found. In the agariceæ the different

kinds of vellum distinguish between species, and the color of the

conidia is also of differential importance. In all cases of distinct

characteristic habits of reproduction and form, one or more different

chemical compounds is found.

In the next group of the musiceæ, or mosses, is an absence of some

chemical compounds that were characteristic of the classes just

described. Many of the albuminous substances are present. Starch[14]

is found often in large quantities, and also oily fats, which are

contained in the oil bodies of the liverworts; wax,[15] organic acids,

including aconitic acid, and tannin, which is found for the first time

at this evolutionary stage of the plant kingdom.

The vascular cryptogams are especially characterized by their mineral

composition.[16] The ash is extraordinarily rich in silicic acid and

alumina.

Equisetum[17]..........silicic acid 60 per cent.

Aspidium............... " " 13

Asplenium.............. " " 35

Osmunda................ " " 53

Lycopodium[18]......... " " 14

" ........ alumina 26 to 27

" ........ manganese 2 to 2.5

These various plants contain acids and compounds peculiar to

themselves.

As we ascend in the plant scale, we reach the phanerogams. These

plants are characterized by the production of true seeds, and many

chemical compounds not found in lower plants.

It will be convenient in speaking of these higher groups to follow M.

Heckel's[19] scheme of plant evolution. All these plants are grouped

under three main divisions: apetalous, monocotyledonous, and

dicotyledonous; and these main divisions are further subdivided.

It will be observed that these three main parallel columns are divided

into three general horizontal planes.

On plane 1 are all plants of simplicity of floral elements, or parts;

for example, the black walnut, with the simple flower contained in a

catkin.

On plane 2 plants which have a multiplicity of floral elements, as the

many petals and stamens of the rose; and finally, the higher plants,

the orchids among the monocotyledons and the composite among the

dicotyledonous plants, come under the third division of condensation

of floral elements.

It will be impossible to take up in order for chemical consideration

all these groups, and I shall restrict myself to pointing out the

occurrence of certain constituents.

I desire now to call attention to chemical groups under the apetalous

plants having simplicity of floral elements.

_Cassuarina equisetifolia_[20] possibly contains tannin, since it is

used for curing hides. The bark contains a dye. It is said to resemble

_Equisetum_[21] in appearance, and in this latter plant a yellow dye

is found.

The _Myrica_[22] contains ethereal oil, wax, resin, balsam, in all

parts of the plant. The root contains in addition fats, tannin, and

starch, also myricinic acid.

In the willow and poplar,[23] a crystalline, bitter substance, salicin

or populin, is found. This may be considered as the first appearance

of a real glucoside, if tannin be excluded from the list.

The oak, walnut, beech, alder, and birch contain tannin in large

quantities; in the case of the oak, ten to twelve per cent. Oak galls

yield as much as seventy per cent.[24]

The numerous genera of pine and fir trees are remarkable for ethereal

oil, resin, and camphor.

The plane[25] trees contain caoutchouc and gum; peppers,[26] ethereal

oils, alkaloids, piperin, white resin, and malic acid. _Datisca

cannabina_[27] contains a coloring matter and another substance

peculiar to itself, datiscin, a kind of starch, or allied to the

glucosides.

Upon the same evolutionary plane among the monocotyledons, the dates

and palms[28] contain in large quantities special starches, and this

is in harmony with the principles of the theory. Alkaloids and

glucosides have not yet been discovered in them.

Other monocotyledonous groups with simplicity of floral elements, such

as the typhaceæ, contain large quantities of starch; in the case of

_Typha latifolia_[29] 12.5 per cent., and 1.5 per cent. gum. In the

pollen of this same plant, 2.08 per cent. starch has been found.

Under the dicotyledonous groups, there are no plants with simplicity

of floral elements.

Returning, now, to apetalous plants of multiplicity and simplification

of floral elements, we find that the urticaceæ[30] contain free formic

acid; the hemp[31] contains alkaloids; the hop,[32] ethereal oil and

resin; the rhubarb,[33] crysophonic acid; and the begonias,[34]

chicarin and lapacho dyes. The highest apetalous plants contain

camphors and oils; the highest of the monocotyledons contain a

mucilage and oils; and the highest dicotyledons contain oils and

special acids.

The trees yielding common camphor and borneol are from genera of the

lauraceæ family; also sassafras camphor is from the same family. Small

quantities of stereoptenes are widely distributed through the plant

kingdom.

The gramineæ, or grasses, are especially characterized by the large

quantities of sugar and silica they contain. The ash of the rice hull,

for example, contains ninety eight per cent. silica.

The ranunculaceæ contain many plants which yield alkaloids, as

_Hydrastia canadensis_, or Indian hemp, _Helleborus_, _Delphinum_,

_Aconitum_, and the alkaloid berberine has been obtained from genera

of this family.

The alkaloid[35] furnishing families belong, with few exceptions, to

the dicotyledons. The colchiceæ, from which is obtained veratrine,

form an exception among the monocotyledons. The alkaloids of the

fungus have already been noted.

[36]Among the greater number of plant families, no alkaloids have been

found. In the labiatæ none has been discovered, nor in the compositæ

among the highest plants.

One alkaloid is found in many genera of the loganiaceæ; berberine in

genera of the berberidaceæ, ranunculaceæ, menispermaceæ, rutaceæ,

papaveraceæ, anonaceæ.

Waxes are widely distributed in plants. They occur in quantities in

some closely related families.

Ethereal oils occur in many families, in the bark, root, wood, leaf,

flower, and fruit; particularly in myrtaceæ, laurineæ, cyperaceæ,

crucifereæ, aurantiaceæ, labiatæ, and umbelliferæ.

Resins are found in most of the higher plants. Tropical plants are

richer in resins than those of cold climates.

Chemical resemblance between groups, as indicating morphological

relations, has been well shown. For example: the similarity[37] of the

viscid juices, and a like taste and smell, among cactaceæ and

portulaceæ, indicate a closer relationship between these two orders

than botanical classification would perhaps allow. This fact was

corroborated by the discovery of irritable stamens in _Portulaca_ and

_Opuntia_, and other genera of cactaceæ.

Darwin[38] states that in the compositæ the ray florets are more

poisonous than the disk florets, in the ratio of about 3 to 2.

Comparing the cycadeæ and palmæ, the former are differently placed by

different botanists, but the general resemblance is remarkable, and

they both yield sago.

Chemical constituents of plants are found in varying quantities during

stated periods of the year. Certain compounds present at one stage of

growth are absent at another. Many facts could be brought forward to

show the different chemical composition of plants in different stages

of growth. The _Thuja occidentalis_[39] in the juvenescent and adult

form, offers an example where morphological and chemical differences

go hand in hand. Analyses of this plant under both conditions show a

striking difference.

Different parts of plants may contain distinct chemical compounds, and

the comparative chemical study of plant orders comprises the analysis

of all parts of plants of different species.

For example; four portions of the _Yucca angustifolia_[40] were

examined chemically; the bark and wood of the root and the base and

blades of the leaves. Fixed oils were separated from each part. These

were not identical; two were fluid at ordinary temperature, and two

were solid. Their melting and solidifying points were not the same.

This difference in the physical character and chemical reaction of

these fixed oils may be due to the presence of free fatty acid and

glycerides in varying proportions in the four parts of the plants. It

is of interest to note that, in the subterranean part of the _Yucca_,

the oil extracted from the bark is solid at the ordinary temperature;

from the wood it was of a less solid consistency; while the yellow

base of the leaf contained an oil quite soft, and in the green leaf

the oil is almost fluid.

Two new resins were extracted from the yellow and green parts of the

leaf. It was proposed to name them _yuccal_ and _pyrophæal_ An

examination of the contents of each extract showed a different

quantitative and qualitative result.

Saponin was found in all parts of the plant.

Many of the above facts have been collected from the investigations of

others. I have introduced these statements, selected from a mass of

material, as evidences in favor of the view stated at the beginning of

this paper.[41] My own study has been directed toward the discovery of

saponin in those plants where it was presumably to be found. The

practical use of this theory in plant analysis will lead the chemists

at once to a search for those compounds which morphology shows are

probably present.

I have discovered saponin in all parts of the _Yucca angustifolia_, in

the _Y. filimentosa_ and _Y. gloriosa_, in several species of agavæ,

and in plants belonging to the leguminosæ family.

The list[42] of plants in which saponin has been discovered is given

in the note. All these plants are contained in the middle plane of

Heckel's scheme. No plants containing saponin have been found among

apetalous groups. No plants have been found containing saponin among

the lower monocotyledons.

The plane of saponin passes from the liliaceæ and allied groups to the

rosales and higher dicotyledons.

Saponin belongs to a class of substances called glucosides. Under the

action of dilute acids, it is split up into two substances, glucose

and sopogenin. The chemical nature of this substance is not thoroughly

understood. The commercial[43] product is probably a mixture of

several substances.

This complexity of chemical composition of saponin is admirably

adapted for the nutrition of the plant, and it is associated with the

corresponding complexity of the morphological elements of the plant's

organs. According to M. Perrey,[44] it seems that the power of a plant

to direct the distribution of its carbon, hydrogen, and oxygen to form

complex glucosides is indicative of its higher functions and

developments.

The solvent action of saponin on resins has been already discussed.

Saponin likewise acts as a solvent upon barium[45] sulphate and

calcium[46] oxalate, and as a solvent of insoluble or slightly soluble

salts would assist the plant in obtaining food, otherwise difficult of

access.

The botanical classifications based upon morphology are so frequently

Saponin is found in endogens and exogens. The line dividing these two

groups is not always clearly defined. Statements pointing to this are

found in the works of Haeckel, Bentham, and others.

Smilax belongs to a transition class, partaking somewhat of the nature

of endogen and of exogen. It is worthy of note that this intermediate

group of the sarsaparillas should contain saponin.

It is a significant fact that all the groups above named containing

saponin belong to Heckel's middle division.

It may be suggested that saponin is thus a constructive element in

developing the plant from the multiplicity of floral elements to the

cephalization of those organs.

It has been observed that the composite occurs where the materials for

growth are supplied in greatest abundance, and the more simple forms

arise where sources of nutrition are remote. We may gather from this

fact that the simpler organs of plants low in the evolutionary scale

contain simpler non-nitrogenous chemical compounds for their

nutrition.

The presence of saponin seems essential to the life of the plant where

it is found, and it is an indispensable principle in the progression

of certain lines of plants, passing from their lower to their higher

stages.

Saponin is invariably absent where the floral elements are simple; it

is invariably absent where the floral elements are condensed to their

greatest extent. Its position is plainly that of a factor in the great

middle realm of vegetable life, where the elements of the individual

are striving to condense, and thus increase their physiological action

and the economy of parts.

It may be suggested as a line of research to study what are the

conditions which control the synthesis and gradual formation of

saponin in plants. The simpler compounds of which this complex

substance is built up, if located as compounds of lower plants, would

indicate the lines of progression from the lower to the saponin

groups.

In my paper[47] read in Buffalo at the last meeting of the American

Association for the Advancement of Science, various suggestions were

offered why chemical compounds should be used as a means of botanical

classification.

The botanical classifications based upon morphology are so frequently

unsatisfactory, that efforts in some directions have been made to

introduce other methods.[48]

There has been comparatively little study of the chemical principles

of plants from a purely botanical view. It promises to become a new

field of research.

The leguminosæ are conspicuous as furnishing us with important dyes,

e.g., indigo, logwood, catechin. The former is obtained principally

from different species of the genus _Indigofera_, and logwood from the

_Hæmatoxylon_ and _Saraca indica_.

The discovery[49] of hæmatoxylin in the _Saraca indica_ illustrates

very well how this plant in its chemical, as well as botanical,

character is related to the _Hæmatoxylon campechianum_; also, I found

a substance like catechin in the _Saraca_. This compound is found in

the _acacias_, to which class _Saraca_ is related by its chemical

position, as well as botanically. Saponin is found in both of these

plants, as well as in many other plants of the leguminosæ. The

leguminosæ come under the middle plane or multiplicity of floral

elements, and the presence of saponin in these plants was to be

expected.

From many of the facts above stated, it may be inferred that the

chemical compounds of plants do not occur at random. Each stage of

growth and development has its own particular chemistry.

It is said that many of the constituents found in plants are the

result of destructive metabolism, and are of no further use in the

plant's economy. This subject is by no means settled, and even should

we be forced to accept that ground, it is a significant fact that

certain cells, tissues, or organs peculiar to a plant secrete or

excrete chemical compounds peculiar to them, which are to be found in

one family, or in species closely allied to it.

It is a fact that the chemical compounds are there, no matter why or

whence they came. They will serve our purposes of study and

classification.

The result of experiment shows that the presence of certain compounds

is essential to the vigor and development of all plants and particular

compounds to the development of certain plants. Plant chemistry and

morphology are related. Future investigations will demonstrate this

relation.

In general terms, we may say that amides and carbohydrates are

utilized in the manufacture of proteids. Organic acids cause a

turgescence of cells. Glucosides may be a form of reserve food

material.

Resins and waxes may serve only as protection to the surfaces of

plants; coloring matters, as screens to shut off or admit certain of

the sun's rays; but we are still far from penetrating the mystery of

life.

A simple plant does what animals more highly endowed cannot do. From

simplest substances they manufacture the most complex. We owe our

existence to plants, as they do theirs to the air and soil.

The elements carbon, oxygen, hydrogen, and nitrogen pass through a

cycle of changes from simple inorganic substances to the complex

compounds of the living cell. Upon the decomposition of these bodies

the elements return to their original state. During this transition

those properties of protoplasm which were mentioned at the beginning,

in turn, follow their path. From germination to death this course

appears like a crescent, the other half of the circle closed from

view. Where chemistry begins and ends it is difficult to say.--_Jour.

Fr. Inst._

[Footnote 1: A lecture delivered before the Franklin Institute,

January 24, 1887.]

[Footnote 2: Studien uber das Protoplasm, 1881.]

[Footnote 3: Vines, p. 1. Rostafinski: Mem. de la Soc. des Sc.

Nat. de Cherbourg, 1875. Strasburger: Zeitschr., xii, 1878.]

[Footnote 4: Botany: Prantl and Vines. London, 1886, p. 110.]

[Footnote 5: For the literature of starch, see p. 115, Die

Pflanzenstoffe, von Hilger and Husemann.]

[Footnote 6: Kutzing: Arch. Pharm., xli, 38. Kraus and Millardet:

Bul. Soc. Sciences Nat., Strasbourg, 1868, 22. Sorby: Jour. Lin.

Soc., xv, 34. J. Reinke: Jahrb. Wissenscht. Botan., x, B. 399.

Phipson: Phar. Jour. Trans., clxii, 479.]

[Footnote 7: Prantl and Vines, p. 111.]

[Footnote 8: L. Crie: Compt. Rend., lxxxviii, 759 and 985. J. De

Seynes, 820, 1043.]

[Footnote 9: Page 279.]

[Footnote 10: M. Nencki and F. Schaffer. N. Sieher: Jour. Pract.

Chem., 23, 412.]

[Footnote 11: E. Klein: Quar. Jour. Micros. Science, 1875, 381. O.

Helm: Arch. Pharm., 1875, 19-24. G. Gugini: Gaz. Chem., 7, 4. W.

Thorner: Bul. Ber, xi, 533.]

[Footnote 12: Handbook of Dyeing. By W. Crookes, London, 1874. p. 367.

Schunck: Ann. Chem. Pharm., 41, 157; 54, 261; 61, 72; 61, 64; 61, 78.

Rochelder and Heldt, ibid., 48, 2; 48, 9. Stenhouse, ibid., 68, 57;

68, 72; 68, 97, 104; 125, 353. See also researches of Strecker, O.

Hesse, Reymann, Liebermann, Lamparter, Knop, and Schnedermann.]

[Footnote 13: Stahlschmidt.]

[Footnote 14: E. Treffner: Inaugur. Diss. Dorpat, 1880.]

[Footnote 15: W. Pfeffer: Flora, 1874.]

[Footnote 16: Die Pflanzenstoffe, p. 323 W. Lange: Bul. Ber., xi,

822.]

[Footnote 17: Ann. Chim. Phys., 41, 62, 208; Ann. Chim. Pharm.,

77, 295.]

[Footnote 18: Fluckiger: Pharmakognosie. Kamp: Ann. Chim. Pharm.,

100, 300.]

[Footnote 19: Revue Scientifiqe, 13 Mars, 1886.]

[Footnote 20: Dictionary of Economic Plants. By J. Smith. London,

1882, p. 294.]

[Footnote 21: Ibid., p. 160. Pharmakognosie des Pflanzenreichs,

Wittstein, p. 736. Ann. Chem. Pharm., 77, 295.]

[Footnote 22: Rabenhorst: Repert. Pharm., lx, 214. Moore: Chem.

Centralbl., 1862, 779, Dana.]

[Footnote 23: Johansen: Arch. Pharm., 3, ix, 210. Ibid., 3, ix

103. Bente: Berl. Ber., viii, 476. Braconnot: Ann. Chim. Phys., 2,

44, 296.]

[Footnote 24: Wittstein; Pharm. des Pflanzenreichs, p. 249.]

[Footnote 25: John; Ibid., p. 651.]

[Footnote 26: Dulong. Oersted, Lucas, Pontet; Ibid., p. 640.]

[Footnote 27: Braconnot: Ann. Chim. Phys., 2, 3. 277. Stenhouse:

Ann. Chim. Phann., 198, 166].

[Footnote 28: 3 Pflanzenstoffe, p. 412.]

[Footnote 29: Lecocq: Braconnot: Pharmacog. Pflan, p. 693.]

[Footnote 30: Gorup-Besanez.]

[Footnote 31: Siebold and Brodbury: Phar. Jour. Trans., 3, 590,

1881, 326.]

[Footnote 32: Wagner: Jour. Prakt. Chem., 58, 352. B. Peters, v.

Gohren: Jahresb. Agric., viii, 114; ix, 105; v. 58. Ann. Jour.

Pharm., 4, 49.]

[Footnote 33: Dragendorff: Pharm. Zeitschr. Russ., xvii, 65-97.]

[Footnote 34: Bonssingault: Ann. Chim. Phys., 2, 27, 315. Erdmann:

Jour. Pract. Chem., 71, 198.]

[Footnote 35: Die Pflanzenstoffe, p. 21.]

[Footnote 36: Ibid.]

[Footnote 37: Meehan: Proc. Acad. Nat. Sciences.]

[Footnote 38: Different forms of flowers on plants of the same

species. Introduction.]

[Footnote 39: Meehan: Proc. Acad. Nat. Sciences.]

[Footnote 40: H.C. De S. Abbott: Trans. Amer. Philos. Soc., 1886.]

[Footnote 41: For further facts confirming this theory, see

"Comparative Chemistry of Higher and Lower Plants." By H.C. De S.

Abbott. Amer. Naturalist, August, 1887.]

[Footnote 42: Different genera and species of the following:

Ranunculaceæ, Berberidaceæ, Carophyllaceæ, Polygalaceæ,

Bromeliaceæ, Liliaceæ, Smilaceæ, Yuccas, Amaryllideæ, Leguminosæ,

Primulaceæ, Rosaceæ, Sapindaceæ, Sapotaceæ]

[Footnote 43: Kobert: Chem Ztg.]

[Footnote 44: Compt. Rend., xciv, p. 1124.]

[Footnote 45: Bul. de la Soc. Chim.]

[Footnote 46: "Yucca angus." Trans. Am. Philos. Soc., Dec., 1885.]

[Footnote 47: Botanical Gazette, October, 1886.]

[Footnote 48: Borodin: Pharm. Jour. Trans., xvi, 369. Pax. Firemy:

Ann. Sci. Nat., xiii.]

[Footnote 49: H.C. De S. Abbott, Proc. Acad. Nat. Sciences, Nov.

30, 1886.]

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