are incompatible with the theory of evolution. I have, therefore,
consistently resisted them for thirty-eight years, and consider them
to be so dangerous that I will briefly give my reasons. First, let
us clearly understand the modern definition of the cell. It is now
generally defined (in accordance with the second thesis) as being
composed of two essentially different parts, the nucleus and the
cell-body, and it is added that these organella differ constantly
both in respect of chemistry, morphology, and physiology. If that
is really so, the cell cannot possibly be the primitive organism; if
it were, we should have a miracle at the beginning of organic life
on the earth. The theory of natural evolution clearly and distinctly
demands that the cell (in this sense) is a secondary development from
a simpler, primary, elementary organism, a homogeneous cytode. There
are still living to-day very simple protists which do not tally with
this definition, and which I designated _monera_ in 1866. As they must
necessarily have preceded the real cells, they may also be called
"precellular organisms."
The earliest organisms to live on the earth, with which the wonderful
drama of life began, can, in the present condition of biological
science, only be conceived as homogeneous particles of plasm--biogens
or groups of biogens, in which there was not yet the division of
nucleus and cell-body which characterizes the real cell. I gave the
name "cytodes" to these unnucleated cells in 1866, and joined them
with the real nucleated cells under the general head of "plastids."
I also endeavored to prove that such cytodes still exist in the form
of independent monera, and in 1870 I described in my _Monograph on
the Monera_ a number of protists which do not tally with the above
definition.
Fifty years ago I made the first careful observations of living monera
(_protamœba_ and _protogenes_), and described them in my _General
Morphology_ (vol. i., pp. 133-5; vol. ii., p. xxii.) as structureless
organisms without organs and the real beginnings of organic life.
Soon afterwards, during a stay in the Canary Islands, I succeeded
in following the continuous life-history of a related organism of
the rhizopod type, which behaved like a very simple mycetozoon, but
differed in having no nucleus; I have reproduced the picture of it in
the first plate of my _History of Creation_. The description of this
orange-red globule of plasm (_protomyxa aurantiaca_) appeared first in
my _Monograph on the Monera_. Most of the organisms which I comprised
under this name exhibited the same movements as true rhizopods (or
sarcodina). It was afterwards proved of some of them that there was
a nucleus hidden within the homogeneous particle of plasm, and that,
therefore, they must be regarded as real cells. But this discovery
was wrongly extended to the whole of the monera, and the existence of
unnucleated organisms was denied altogether. Nevertheless, there are
living to-day several kinds of these organisms without organs, some
of them being very widely distributed. The chief examples are the
chromacea and the bacteria, the former with vegetal and the latter with
animal metabolism (or the former plasmodomous = plasma-forming, and the
latter plasmophagous = plasma-feeding). On the ground of this important
chemical difference, I distinguished two principal groups of the monera
in my _Systematic Phylogeny_ twenty years ago--the phytomonera and
the zoomonera, the former being unnucleated protophyta and the latter
unnucleated protozoa.
Among living organisms the chromacea are certainly the most primitive
and the nearest to the oldest inhabitants of the earth. Their simplest
forms, the chroococcacea, are nothing but small structureless particles
of plasm, growing by plasmodomism (formation of plasm) and multiplying
by simple cleavage as soon as their growth passes a certain limit
of individual size. Many of them are surrounded by a thin membrane
or somewhat thicker gelatinous covering, and this circumstance had
prevented me for some time from counting the chromacea as monera.
However, I became convinced afterwards that the formation of a
protective cover of this kind around the homogeneous particle of
plasm may indeed be regarded from the physiological stand-point as a
"purposive" structure, but at the same time may be looked upon, from
the purely physical stand-point, as a result of superficial strain.
On the other hand, the physiological character of these plasmodomous
monera is especially important, as it gives us the simple key to the
solution of the great question of spontaneous generation (or archigony,
_cf._ chapter xv.).
The chromacea are to-day found in every part of the earth, living
sometimes in fresh water and sometimes in the sea. Many species form
blue-green, violet, or reddish deposits on rocks, stones, wood, and
other objects. In these thin gelatinous plates millions of small
homogeneous cytodes are packed close together. Their tint is due to a
peculiar coloring matter (phycocyan), which is chemically connected
with the substance of the plasma-particle. The shade of this color
differs a good deal in the various species of chromacea (of which more
than eight hundred have been distinguished); in the native species it
is generally blue-green or sage-green, sometimes blue, cyanine blue,
or violet. Hence the common name cyanophyceæ (_i.e._, blue algæ).
It is incorrect, for two reasons; firstly, because only a part of
these protophyta are blue, and, secondly, because they (as simple,
primitive plants without tissue) must be distinguished from the real
algæ (phyceæ), which are multicellular, tissue-forming plants. Other
chromacea are red, orange, or yellow in color, as the interesting
_trichodesmium erythræum_, for instance, the flaky masses of which,
gathering in enormous quantities, cause at certain times the yellow
or red coloring of the sea-water in the tropics; it is these that are
responsible for the name "Red Sea" on the Arabian and "Yellow Sea" on
the Chinese coast. When I passed the equator in the Sunda Straits on
March 10, 1901, the boat sailed through colossal accumulations, several
miles in width, of this trichodesmium. The yellow or reddish surface of
the water looked as if it were strewn with sawdust. In the same way,
the surface of the Arctic Ocean is often colored brown or reddish-brown
by masses of the brown _procytella primordialis_ (formerly described as
_protococcus marinus_).
It is clearly quite illogical to regard the chromacea as a class
or family of the algæ, as is still done in most manuals of botany.
The real algæ--excluding the unicellular diatomes and paulotomes,
which belong to the protophyta--are multicellular plants that form
a _thallus_ or bed of a certain form and characteristic tissue. The
chromacea, which have not advanced as far as the real nucleated cell,
are unnucleated cytodes of a lower and earlier stage of plant-life.
If one would compare the chromacea with algæ or other plants at all,
the comparison cannot be with their constituent cells, but merely
with the chromatophora or chromatella, which are found in all green
plant-cells, and form _part_ of their contents. To be more precise,
these green granules of chlorophyll must be regarded as organella of
the plant-cell, or separated plasma-formations which arise beside the
nucleus in the cytoplasm. In the embryonic cells of the germs of plants
and in their vegetation points the chromatophora are as yet colorless,
and are developed, as solid, very refractive, globular, or roundish
granules, from the firm layer of plasm which immediately surrounds the
nucleus. Afterwards they are converted, by a chemical process, into
the green chlorophyll granules or chloroplasts, which have the most
important function in the plasmodomism or carbon-assimilation of the
plant.
The fact that the green chlorophyll granules grow independently
within the living plant-cell and multiply by segmentation is very
important and interesting. The globular chloroplasts are constricted
in the middle, and split into two equal daughter-globules. These
daughter-plastids grow, and multiply in turn in the same way. Hence
they behave within the plant-cell just like the free-living chromacea
in the water. On the strength of this significant comparison, one of
our ablest and most open-minded scientists, Fritz Müller-Desterro, of
Brazil, pointed out in 1893 that we may see in every green vegetal cell
a symbiosis between plasmodomous green and plasmophagous not-green
companions (_cf._ my _Anthropogeny_, figs. 277 and 278, and in the
text).
Many species of the simplest chromacea live as monobia (individually).
When the tiny plasma globules have split into two equal halves by
simple segmentation, they separate, and live their lives apart. This
is the case with the common, ubiquitous chroococcus. However, most
species live in common, the plasma granules forming more or less thick
cœnobia, or communities or colonies of cells. In the simplest case
(_aphanocapsa_) the social cytodes secrete a structureless gelatinous
mass, in which numbers of blue-green plasma globules are irregularly
distributed. In the _glœocapsa_, which forms a thin blue-green
gelatinous deposit on damp walls and rocks, the constituent cytodes
cover themselves immediately after cleavage with a fresh gelatinous
envelope, and these run together into large masses. But the majority
of the chromacea form firm, threadlike cell communities or chains
of plastids (catenal cœnobia.) As the transverse cleavage of the
rapidly multiplying cytodes always follows the same direction, and
the new daughter-cytodes remain joined at the cleavage surfaces, and
are flattened into discoid shape, we get stringlike formations or
articulated threads of considerable length, as in the oscillaria and
nostochina. When a number of these threads are joined together in
gelatinous masses, we often get large, irregular, jelly-like bodies, as
in the common "shooting-star jellies" (_nostoc communis_). They attain
the size of a plum.
In view of the extreme importance which I attach to the chromacea as
the earliest and simplest of all organisms, it is necessary to put
clearly the following facts with regard to their anatomic structure and
physiological activity:
1. The organism of the simplest chromacea is _not_ composed of
different organella or organs; and it shows no trace of purposive
construction or definite architecture.
2. The homogeneous tinted plasma granule which makes up the entire
organism in the simplest case (_chroococcus_) exhibits no plasma
structure (honeycomb, threads, etc.) whatever.
3. The original globular form of the plasma particle is the simplest of
all fundamental types, and is also that assumed by the inorganic body
(such as a drop of rain) in a condition of stable equilibrium.
4. The formation of a thin membrane at the surface of the structureless
plasma granule may be explained as a purely physical process--that of
surface strain.
5. The gelatinous envelope which is secreted by many of the chromacea
is also formed by a simple physical (or chemical) process.
6. The sole essential vital function that is common to all the
chromacea is self-maintenance, and growth by means of their vegetal
metabolism, or plasmodomism (=carbon assimilation); this purely
chemical process is on a level with the catalysis of inorganic
compounds (chapter x.).
7. The growth of the cytodes, in virtue of their continuous
plasmodomism, is on a level with the physical process of crystal growth.
8. The reproduction of the chromacea by simple cleavage is merely the
continuation of this simple growth process, when it passes the limit of
individual size.
9. All the other vital phenomena which are to be seen in some of the
chromacea can also be explained by physical or chemical causes on
mechanical principles. Not a single fact compels us to assume a "vital
force."
Especially noteworthy in regard to the physiological character of these
lowest organisms are their bionomic peculiarities, especially the
indifference to external influences, higher and lower temperatures,
etc. Many of the chromacea live in hot springs, with a temperature
of fifty to eighty degrees centigrade, in which no other organism is
found. Other species may remain for a long time frozen in ice, and
resume their vital activity as soon as it thaws. Many chromacea may be
completely dried up, and then resume their life if put in water after
several years.
Next in order to the chromacea we have the bacteria, the remarkable
little organisms which have been well known in the last few decades
as the causes of fatal diseases, and the agents of fermentation,
putrefaction, etc. The special science which is concerned with
them--modern bacteriology--has attained so important a position in
a short period--especially as regards practical and theoretical
medicine--that it is now represented by separate chairs at most of
the universities. We may admire the penetration and the perseverance
with which scientists have succeeded, with the aid of the best modern
microscopes and methods of preparation and coloring, in making so
close a study of the organism of the bacteria, determining their
physiological properties, and explaining their great importance for
organic life by careful experiments and methods of culture. The
bionomic or economic position of the bacteria in nature's household
has thus secured for these tiny organisms the greatest scientific and
practical interest.
However, we find that certain general views have been maintained by
specialists in bacteriology up to our own time which are in curious
contrast with these brilliant results. The biologist who studies the
systematic relations of the bacteria from the modern point of view of
the theory of descent is bewildered at the extraordinary views as to
the place of the bacteria in the plant-world (as segmentation-fungi),
their relations to other classes of plants, and the formation of their
species. When we carefully consider the morphological properties that
are common to all true bacteria and compare them with other organisms,
we are forced to the conclusion that I urged years ago in various
writings: the bacteria are not real (nucleated) cells, but unnucleated
cytodes of the rank of the monera; they are not real (tissue-forming)
fungi, but simple protists; their nearest relatives are the chromacea.
The individual organisms of the simplest kind, which bacteriologists
call "bacteria-cells," are not real nucleated cells. That is the clear
negative result of a number of most careful investigations which
have been made up to date with the object of finding a nucleus in
the plasma-body of the bacteria. Among recent exact investigations
we must especially note those of the botanist Reinke, of Kiel, who
sought in vain to detect a nucleus in one of the largest and most
easily studied genera of the bacteria, the _beggiatoa_, using every
modern technical aid. His conviction that this important cell-structure
is really lacking is the more valuable, as it is very prejudicial
to his own theory of "dominants." Other scientists (especially
Schaudinn) have recently claimed, as equivalent to a nucleus in some
of the larger bacteria, a number of very small granules, which are
irregularly distributed in the plasm, and are strongly tinted under
certain coloring processes. But even if the chemical identity of these
substances which take the same color were proved--which is certainly
not the case--and even if the appearance of scattered nuclein-granules
in the plasm could be regarded as a preliminary to, or a beginning
of, the differentiation of an individual, morphologically distinct
nucleus, we should not yet have shown its independence as an organellum
of the cell.
Nor is this any more proved from the circumstance that in some bacteria
(not all) we find a severance of the plasm into an inner and outer
layer, or a frothy structure with vacuole-formation, or a special
sharply outlined membrane on the plastid. Many bacteria (but not all)
have such a membrane, like the nearly related chromacea, and also the
secretion of a gelatine envelope. Both classes have also in common
an exclusively monogenetic reproduction. The bacteria multiply, like
the chromacea, by simple segmentation; as soon as the structureless
plasma-granule has reached a certain size by simple growth, it is
constricted and splits into two halves. In the long-bodied bacteria
(the rod-shaped bacilli) the constriction always goes through the
middle of the long axis, and is, therefore, simple transverse cleavage.
Many bacteria have also been said to multiply by the formation of
spores. But these so-called "spores" are really permanent quiescent
forms (without any multiplication of individuals); the central part
of the plastid (endoplasm) condenses, separates from the peripheral
part (exoplasm), and undergoes a chemical change which makes it very
indifferent to external influences (such as a high temperature).
The great majority of the bacteria differ so little morphologically
from the chromacea that we can only distinguish these two classes
of monera by the difference in their metabolism. The chromacea,
as protophyta, are plasmodomous. They form new plasm by synthesis
and reduction from simple inorganic compounds--water, carbonic
acid, ammonia, nitric acid, etc. But the bacteria, as protozoa, are
plasmophagous. They cannot, as a rule, form new plasm, but have to
take it from other organisms (as parasites, saprophytes, etc.);
they decompose it by analysis and oxydation. Hence the colorless
bacteria are without the important green, blue, or red coloring matter
(phycocyan) which tints the plastids of the chromacea, and is the real
instrument of the carbon-assimilation. However, there are exceptions
in this respect: _bacillus virens_ is tinted green with chlorophyll,
_micrococcus prodigiosus_ is blood-red, other bacteria purple, and
so on. Certain earth-dwelling bacteria (_nitro-bacteria_) have the
vegetal property of plasmodomism; they convert ammonia by oxydation
into nitrous acid, and this into nitric acid, using as their source of
carbon the carbonic acid gas in the atmosphere. They are thus quite
independent of organic substances, and feed, like the chromacea, on
simple inorganic compounds.
Hence the affinity between the plasmodomous chromacea and plasmophagous
bacteria is so close that it is impossible to give a single safe
criterion that will effectually separate the two classes. Many
botanists accordingly combine both groups in a single class with the
name of _schizophyta_, and within this distinguish as "orders" the
blue-green chromacea as _schizophycæ_ (cleavage-algæ) and the colorless
bacteria as _schizomycetes_ (cleavage-fungi). However, we must not
take this division too rigidly; and the absolute lack of a nucleus
and tissue-formation separates the chromacea just as widely from the
multicellular tissue-forming algæ as the bacteria from the fungi. The
simple multiplication by the halving of the cell, which is expressed in
the name "cleavage-plants" (_schizophyta_), is also found in many other
protists.
The number of forms that can be distinguished as species in the
technical sense is very great in the case of the bacteria, in
spite of the extreme simplicity of their outward appearance; many
biologists speak of several hundred, and even of more than a thousand,
species. But when we look solely to the outer form of the living
plasma-granule, we can only distinguish three fundamental types: (1)
Micrococci, or spherobacteria (briefly, cocci), globular or ellipsoid;
(2) bacilli, or rhabdo-bacteria (also called eubacteria, or bacteria in
the narrower sense), rod-shaped, cylindrical, and often twisted like
worms (comma-bacilli); (3) spirilla, or spirobacteria, screw-shaped
rods (vibriones when the screw is slight, and spirochæta when it
has many coils). Besides this threefold difference in the forms of
the cytodes, we have a ground of distinction in many bacilli and
spirilla in the possession of one or more very thin lashes (flagella),
which proceed from one of both poles of the lengthened plastid. The
construction and vibration of these serves for locomotion in the
swimming bacteria; but they are only found for a time in many species,
and in many others are altogether wanting.
Since, then, neither the simple outer form of the bacterium-cytodes
nor their homogeneous internal structure provides a satisfactory
ground for the systematic distinction of the numerous species,
their physiological properties are generally used for the purpose,
especially their different behavior towards organic foods (albumin,
gelatine, etc.), their chemical actions, and the various effects of
poisoning and decomposition which they produce in the living organism.
No bacteriologist now doubts that all the vital activities of the
bacteria are of a chemical nature, and precisely on this account
these microbes are of extreme importance. When we bear in mind how
complicated are the relations of the various species of bacteria to
the tissues of the human body, in which they cause the diseases of
typhus, hypochondriasis, cholera, and tuberculosis, we are bound to
admit that the real cause of these maladies must be sought in the
peculiar molecular structure of the bacterium-plasm, or the particular
arrangement of its molecules and the innumerable atoms (more than a
thousand) which are, in a very loose way, made up into special groups
of molecules. The chemical products of their mutual action are what we
call ptomaines, which are partly very virulent poisons (toxins). We
have succeeded in producing several of these poisonous matters in large
quantities by artificial culture, and isolating them and experimentally
ascertaining their nature; as, for instance, tetanin, which causes
tetanus, typhotoxin, the poison of typhus, etc.
In thus declaring the action of bacteria to be purely chemical and
analogous to that of well-known inorganic poisons, I would particularly
point out that this very justifiable statement is a pure hypothesis;
it is an excellent illustration of the fact that we cannot get on
in the explanation of the most important natural phenomena without
hypotheses. We can see nothing whatever of the chemical molecular
structure of the plasm, even under the highest power of the microscope;
it lies far below the limit of microscopic perception. Nevertheless,
no expert scientist has the slightest doubt of its existence, or that
the complicated movements of the sensitive atoms and the molecules and
groups of molecules they make up are the causes of the vast changes
which these tiny organisms effect in the tissues of the human and the
higher animal body.
Moreover, the distinction of the many species of bacteria is of
interest in connection with the general question of the nature and
constancy of a species. Whereas formerly in biological classification
only definite morphological characters, or definable differences in
outer form or inner structure, were regarded as of any moment in the
distinction of species, here, in view of the vagueness or total lack
of these characters, we have to look mainly to the physiological
properties, and these are based on the chemical differences in their
hypothetical molecular structure. But even these are not absolutely
constant; on the contrary, many bacteria lose their specific qualities
by progressive culture under changed food-conditions. By a change in
the temperature and the nutritive field in which a number of poisonous
bacteria have been reared, or by the action of certain chemicals, not
only the growth and multiplication are altered, but also the injurious
effect they have on other organisms by the generation of poisons.
This poisonous effect is weakened, and--what is most important--the
weakening is transmitted by heredity to the following generations. On
this is based the familiar process of inoculation, an admirable example
of the inheritance of acquired characteristics.
As the bacteria are still often described as "cleavage-fungi" and
classified along with the real fungi, we must particularly point
out the wide gulf that separates the two groups. The real fungi (or
_mycetes_) are metaphyta, their multicellular body (_thallus_) forming
a very characteristic sort of tissue, the mycelium; this is composed
of a number of interlaced and interwoven threads (or hyphens). Each
fungus-thread consists of a row of lengthened cells, which have a thin
membrane and enclose a number of small nuclei in the colorless plasm.
Moreover, the two sub-classes of the real fungi, the ascomycetes and
basimycetes, form peculiar fruit-bodies which generate spores (ascodia
and basidia). There is no trace whatever of these real characteristics
of the true fungus in the bacteria. Nor is it less incorrect to class
them with the fungilli, the so-called unicellular fungi or phycomycetes
(ovomycetes and zygomycetes); these form a special class of protists
which has the closest affinity to the gregarinæ.
Like the closely related chromacea, many of the bacteria show a marked
tendency to form communities or cell-colonies. These cell-communities
arise, as elsewhere, from the fact that the individuals, which
multiply rapidly by continuous cleavage, remain joined together.
This may happen in two ways. When the social bacteria secrete large
quantities of gelatine, and remain distributed in this, we have the
_zooglœa_ (as in the case of the _aphanocapsa_ and _glœocapsa_
among the chromacea). If, on the other hand, the long-bodied
bacilli remain fastened together in rows, we get the knotted
threads of _leptothrix_ and _beggiatoa_ (which may be compared with
the oscillaria). And, if these threads go into branches, we have
_cladothrix_. Other cœnobia of bacteria have the appearance of
disks, the cytodes dividing in a plane, usually in groups of four (as
in _merismopedia_), or of cube-shaped packets when they are in all
three directions of space (_sarcina_).
The two classes of bacteria and chromacea seem, in the present
condition of our knowledge, on account of their simple organization,
to be the simplest of all living things, real monera, or organisms
without organs. Hence we have to put them at the lowest stage of
the protist kingdom, and must regard the difference between them
and the most highly differentiated unicellular beings (such as the
radiolaria, ciliated infusoria, diatomes, or siphonea) as no smaller
than the difference (in the realm of the histona) between a lower
polyp (_hydra_) and a vertebrate, or between a simple alga (_ulva_)
and a palm. But if the kingdom of the protists is badly divided, on
the older rule, into a plant kingdom and an animal kingdom, the only
discriminating mark we have left is the difference in metabolism; in
that case we have to include the plasmophagous bacteria in the animal
kingdom (as Ehrenberg did in 1838) and the plasmodomous chromacea
in the plant kingdom. The remarkable class of the flagellata, which
includes ciliated unicellulars of both groups, contains several
forms which are only distinguished from the typical bacterium by the
possession of a nucleus. If it is true that in some of the protists
which were counted as bacteria a real nucleus has been detected, these
must be separated from the others (unnucleated) and included in the
nucleated flagellata.
The monera which I described in 1866, and on which I based the theory
of the monera in my monograph, belong to a different division of the
protists from the classes of bacteria and chromacea. These are the
forms which I described as _protamœba_, _protogenes_, _protomyxa_,
etc. Their naked mobile plasma-bodies thrust out pseudopodia, or
variable "false feet," from their surface, like the (nucleated) real
rhizopods (=sarcodinæ); but they differ essentially from the latter in
the absence of a nucleus. Afterwards (in my _Systematic Phytogeny_)
I proposed to separate these unnucleated rhizopods from the others,
giving the name of _lobomonera_ (_protamœba_) to the amœba-like
monera with flap-shaped feet, and the name of _rhizomonera_
(_protomyxa_, _pontomyxa_, _biomyxa_, _arachnula_, etc.) to the
gromia-like, root-feet forming monera. However, of late years, real
nuclei have been detected in each of these large monera, and so they
have been proved to be true cells. This discovery was made possible by
the improved modern methods of coloring the nucleus which I had not the
use of thirty years ago in my first observations. On the strength of
these recent discoveries many scientists claim that all the monera I
described are true cells, and must have nuclei. This baseless assertion
is much employed by the opponents of the theory of evolution in order
to deny the existence of the monera altogether.
Of the genus of monera which we call protamœba I have given an
illustration in my _History of Creation_ (tenth edition), which has
been frequently reproduced. Several species (at least two or three)
of this genus still exist, and are distinguished by the shape of
their flap-formation and their method of motion. They resemble
ordinary simple amœbæ, and only differ from these to any extent
in the absence of a nucleus. The _protamœba primitiva_ seems
to be pretty widely distributed; it has been found repeatedly by
observers (Gruber, Cienkowski, Leidy, etc.) in inland waters. In the
zoological demonstrations which I have given at the University of Jena
for forty years, and in the course of which the lowly inhabitants
of our fresh water are regularly examined with the microscope, the
_protamœba primitiva_ has been found four or five times. It
always had the same form, as I described it, moved about by the slow
formation of flaps at its surface, multiplied by simple cleavage,
and showed no trace of a nucleus in its homogeneous plasma-body even
with the most careful application of the modern methods of tinting
the nucleus. A larger number of very fine granules (microsoma) that
were irregularly distributed in the plasm, and were more or less
colored by nucleus-reagents, cannot be reckoned as clear equivalents
of the nucleus in this or in similar cases; they are probably products
of metabolism. The same may be said of the larger marine form of
rhizomoneron, which A. Gruber has recently called _pelomyxa pallida_.
The large marine form of rhizomoneron to which Huxley gave the name
of _bathybius Haeckelii_ in 1868, and as to the real nature of which
many opinions have been expressed, seems, according to the latest
investigation, not to have the significance ascribed to it. However,
the much-discussed question of the bathybius is superfluous as far as
our monera theory and the associated hypothesis of archigony (chapter
xv.) are concerned, since we have now a better knowledge of the much
more important monera-forms of the chromacea and bacteria.
In the case of some of the protists I described in my _Monograph on the
Monera_, it is at present doubtful whether their plasma-body contains
a nucleus or not, and, therefore, whether they are to be classed as
true cells or cytodes. This applies especially to the forms which
only happened to come under observation once, such as _protomyxa_
and _myxastrum_. In these obscure cases we must wait for fresh
investigations and the application of the modern methods of tinting
the nucleus. I may, however, point out, in passing, that these famous
methods of nucleus-coloring give by no means the absolute certainty
which is ascribed to them; there are other substances which take color
in the same way as chromatin. As far as my monera theory is concerned,
or the great general importance which I attach to these unnucleated
living granules of plasm, it does not matter whether a nucleus is
detected in these problematic monera or not. The chromacea alone--the
most important of all monera--completely suffice to provide a base for
the far-reaching theoretical conclusions which I draw from it.
At the close of these observations on the monera I will briefly
recapitulate the weighty inferences which we can deduce from their
simple organization. They serve as a solid foundation for the chief
theses of our monistic biology; and they are inconsistent with the
dualistic views of modern vitalists. In the first place, I emphasize
the fact that the structureless plasm-body of the simple monera has
no sort of organization and no composition from dissimilar parts
co-operating for definite vital aims. Reinke's conscious "dominant"--as
well as Weismann's mechanical "determinants"--have nothing to do here.
The whole vital activity of the simplest monera, especially of the
chromacea, is confined to their metabolism, and is therefore a purely
chemical process, that may be compared to the catalysis of inorganic
compounds. The simple formation of individuals in this primitive living
matter is merely a question of the cleavage of plasma globules of a
certain size (_chroococcus_); and their primitive multiplication (by
simple self-division) is only a continued growth (analogous to that
of the crystal). When this simple growth passes a certain limit, that
is fixed by the chemical constitution, it leads to the independent
existence of the redundant growth-products.
X
NUTRITION
Functions of nutrition--Assimilation and disassimilation--Plasmodoma
and plasmophaga--Phytoplasm and zooplasm--Plasmodomism of
plants--Chlorophyll granules and nitro-bacteria--Plasmophagism of
fungi and animals--Metasitism (conversion of metabolism)--Nutrition
of the monera (chromacea, bacteria, rhizomonera)--Nutrition of the
protophyta and metaphyta (cell-plants and tissue-plants)--Nutrition
of the metazoa--- Gastræa theory--Gastro-canal system of the
cœlenteria (gastræads, sponges, cnidaria, platodes)--Nutrition
of the cœlomaria (digestion, circulation, respiration,
evacuation)--Saprositism--Parasitism--Symbiosis.
The wonder of life which we call, in the widest sense of the word,
"nutrition" is the chief factor in the self-maintenance of the organic
individual. It is always bound up with a chemical modification of the
living matter, an organic metabolism (circulation of matter), and a
corresponding circulation of force. In this chemical process plasm is
used up, built up afresh, and once more disintegrated. The metabolism
which lies at the root of this chemistry of food is the essential
feature in the manifold processes of nutrition. A large part of the
several nutritive processes are explained without further trouble
by the known physical and chemical properties of inorganic bodies;
for another part of them we have not yet succeeded in doing this.
Nevertheless, all impartial physiologists now agree that it is possible
in principle, and that we have no reason to introduce a special vital
principle. All the trophic (nutritive) processes, without exception,
are subject to the law of substance.
In all the higher plants and animals the chemical process of
metabolism, with the stream of energy that accompanies it, is a very
complex vital activity, in which many different functions and organs
co-operate with the common aim of self-maintenance. As a rule, they are
distributed in four groups--namely: (1) Intussusception of food and
digestion: (2) distribution of the food in the body, or circulation;
(3) respiration, or exchange of gases; and (4) excretion of unusable
matter. In most of the histona, either tissue-plants or tissue-animals,
a number of organs are differentiated for the accomplishment of these
tasks. At the lower stages of life this division of labor is not found,
the entire process of nutrition being accomplished by a single layer of
cells (lower algæ, gastræads, sponges, lower polyps). In the protists,
again, it is the single cell that does all these things itself; in the
simplest cases, the monera, a homogeneous plasma-globule. As a long
gradation uninterruptedly unites these lowest forms of nutrition with
the more complicated forms, we must regard the latter no less than the
former as physico-chemical processes.
When we take the whole of the metabolic functions in organisms
together, we may look upon them as the outcome of two opposite chemical
processes--on the one hand the building-up of living matter by taking
in food (assimilation), and on the other the breaking-down of it in
consequence of its vital activity (disassimilation). As in every case
the plasm is the active living matter, we may say: _Assimilation_ (or
plasma-production) consists in the conversion within the organism into
the special plasm of the particular species of food that has been
received from without; _disassimilation_ (or plasma-destruction)
is the result of the work done by the plasm, which is the cause of
its partial decomposition or breakdown. In both respects there is a
striking difference between the two great kingdoms of organic nature.
The plant kingdom is, on the whole, the agent of assimilation, forming
new plasm by synthesis and reduction from inorganic matter. In the
animal world, on the contrary, disassimilation preponderates, the plasm
received being resolved by oxydation, and the actual energy taken out
of it by analysis being converted into heat and motion. Plants are
plasmodomous; animals, plasmophagous.
Of all the chemical processes the most important, because the most
indispensable, for the origin and maintenance of organic life is the
constant reconstruction of plasm. We give it the name of plasmodomism
(_domeo_=to build up), or carbon-assimilation. Botanists have the
habit of late of calling it briefly assimilation, and have thus
caused a good deal of misunderstanding. The more common and older
meaning of assimilation in animal physiology is, in the widest sense,
the intussusception and preparation of the food received. But the
carbon-assimilation in plants--what I call plasmodomism--is only the
first and original form of plasma-production. It means that the plant
is able, under the influence of sunlight, to form carbohydrates,
and from these new plasm, out of simple inorganic compounds (water,
carbonic acid, nitric acid, and ammonia) by synthesis and reduction.
The animal is unable to do this. It has to take its plasm in its
food from other organisms--plant-eaters directly, and animal-eaters
indirectly. We therefore give the title of _plasmophagous_ to these
animal "plasma-eaters." In working up the foreign plasm it has eaten,
and converting it into its own specific form of plasm, the animal
also accomplishes assimilation; but this animal albumin-assimilation
is totally different from the vegetal carbon-assimilation. The
fresh-formed animal plasm is then broken up by oxydation, and by this
analysis the energy needed for the vital movements is obtained.
The physiological contrast which we thus find between the two principal
forms of living matter, the synthetic plasm of the plant and the
analytic plasm of the animal, is of great importance for the lasting
maintenance of the whole organic world. It depends on a reversal of
the molecular movement in the plasm, the intimate nature of which is
just as little known to us as the chemical constitution of the albumins
in general, and that of living albumin, the plasm, in particular. As
I mentioned in chapter v., modern physiological chemistry has good
reason to believe that the invisible albumin-molecule is, comparatively
speaking, gigantic, and is composed of more than a thousand atoms.
These are in such an unstable equilibrium, so complicated and
impermanent an arrangement, that the slightest push or stimulus
suffices to alter them and form a new kind of plasm. As a fact, the
number and variety of kinds of plasm are immense. This is seen at
once from the ontogenetic fact that the ovum and sperm-cell of each
species (and each variety) have a specific chemical constitution.
In reproduction this is transmitted to the offspring. But, setting
aside these countless finer modifications, we may distinguish two
chief groups of kinds of plasm: the phytoplasm of the plant, with the
synthetic property of plasmodomism, and the zooplasm of the animal,
which is destitute of this property, and so confined to plasmophagy.
The remarkable synthetic process of building up the plasm, to which
we give the name of plasmodomism, or carbon-assimilation, usually
needs as its first condition the radiant energy of sunlight. Every
green plant-cell contains in its chlorophyll-granules so many tiny
laboratories, their green plasm being able to form new plasm out of
inorganic compounds under the influence of light. The water that is
needed for this, besides nitrogenous compounds (nitric acid, ammonia),
is drawn from the earth by the roots; the carbonic acid is taken from
the atmosphere by the green leaves. The immediate products of the
synthesis, due to the separation of the carbonic acid, is, as a rule, a
non-nitrogenous starch-flour (_amylum_). This is further used for the
composition of the nitrogenous albumin by an as yet unknown synthetic
process, with the aid of nitrogenous mineral compounds. In this process
of reduction the separated free oxygen is returned to the atmosphere.
The carbohydrates that chiefly co-operate in this are glucoses and
maltoses: the mineral substances, especially salts of potassium and
magnesium, and compounds of these elements with nitric acid, sulphuric
acid, and phosphoric acid. Iron is also found to be an important
element in the process, though in a very small quantity. As a rule,
the ferruginous chlorophyll can only form new plasm with the help of
light-waves. The most important part of the spectrum for this purpose
is that containing the red, orange, and yellow waves.
The chief factor in plasma-formation in the organic world is the
photosynthesis, or ordinary carbon-assimilation by chlorophyll, the
wonderful green matter that amounts to only a very small percentage
(about one-tenth) of the weight of the chlorophyll-granules, and can
be separated from their plasmatic substance by certain methods. Even
when the plant has some other color than green the chlorophyll is still
the real plasmodomous substance. Its green color is then masked by
some other color--diatomin in the yellow diatomes, phycorhodin in the
red rhodophyceæ, phycophæin in the brown phæophyceæ, and phyocyan in
the blue-green chromacea or cyanophyceæ. The latter have an especial
interest for us, because in the simplest specimens the entire organism
is merely a globular bluish-green granule of plasm. Moreover, in the
simplest forms of nucleated primitive plants (_algariæ_)--many of the
so-called unicellular algæ--the metabolism is effected by a single
grain of chlorophyll. There is usually a large number of them in the
plasm of the plant-cells.
Another kind of plasm-synthesis, quite different from the ordinary
plasmodomism by chlorophyll and sunlight has lately been discovered in
some of the lowest organisms (by Heraeus, Winogradsky, and others). The
nitro-bacteria (or nitromonades) are tiny monera (unnucleated cells)
that live in complete darkness underground. Their globular colorless
plasma-bodies contain neither chlorophyll nor nucleus. They have the
remarkable capacity of forming carbohydrates, and from these plasm,
by a peculiar synthesis out of purely inorganic compounds--water,
carbonic acid, ammonia, and nitric acid. Pfeffer has called this
carbon-assimilation, on account of its purely chemical nature,
"chemosynthesis," in opposition to the ordinary photosynthesis by
means of sunlight. There are also other bacteria (sulphur-bacteria,
purple-bacteria, etc.) that show various peculiarities of metabolism.
The nitro-bacteria must belong to the oldest monera, and represent a
transition from the vegetal chromacea to the animal bacteria.
The extensive class of the fungi (or _mycetes_) resembles a part of
the bacteria in regard to metabolism. These organisms are, it is true,
generally regarded as plants, but they have not the capacity of the
green, chlorophyll-bearing plants to supply themselves with carbon
from the carbonic acid in the atmosphere. They have to take it from
organic substances, such as albumin, carbohydrates, etc., like the
animals. But while the animals have to derive their nitrogen from the
latter, the fungi can obtain it from inorganic matter in the earth.
Fungi cannot support life without the addition of organic compounds;
but we can make them grow in a food solution consisting of sugar and
purely inorganic nitrogenous salts. Thus they are on the border that
separates the plasmodomous plants from the plasmophagous animals. Like
the latter, the fungi have evolved from the plants through changed food
conditions. We find this process even among the unicellular protists in
the phycomycetes, which descend from the siphonea. In the same way the
real multicellular fungi (ascomycetes and basimycetes) may be traced to
the tissue-forming algæ.
All true animals have to derive their food from the plant kingdom, the
vegetal feeders directly, and the flesh feeders indirectly, when they
consume vegetal feeders. Hence the animals are, in a certain sense, as
the older natural philosophy put it four hundred years ago, "parasites
of the plant world." From the point of view of phylogeny, the animal
kingdom is, therefore, clearly much younger than the plant kingdom. The
development of the animals from the plants was determined originally by
a change in the method of nutrition which we call metasitism.
The chemical modification of the living matter which is connected
with the loss of plasmodomism--in other words, the conversion of the
reducing phytoplasm into oxidizing zooplasm--must be regarded as
one of the most important changes in the history of organic life.
This "reversal of metabolism" is polyphyletic; it has been repeated
many times in the course of biological history, and has taken place
independently in very different groups of the organic world--whenever
a plasmodomous cell or group of cells (=tissue) had occasion to feed
directly on ready-made plasm, instead of giving itself the trouble of
building it up out of inorganic compounds. We see this particularly
among the unicellular protists in the independent ciliated cells.
The longer plasmophagous flagellata, which are colorless, and have
no chlorophyll (monodina, conoflagellata), closely resemble in
form and movement the older plasmadomous and chlorophyll-bearing
mastigota, from which they are descended (volvocina, peridinia); they
only differ in the manner of nutrition. The colorless flagellata
feed on ready-formed plasm, which they obtain either by means of
their lashes or by a special cell mouth in their cell body. On the
other hand, their ancestors, the green or yellow mastigota, form
new plasm by photosynthesis like true cells. But there are also
complete intermediate forms between the two groups--for instance, the
chrysomonades and the gymnodinia; these may behave alternately as
protozoa or protophyta. In the same way we can derive the phycomycetes
by metasitism from the siphonea, the fungi from the algæ; and, finally,
the process is also found in many of the higher parasitic plants
(orchids, orobanches, etc.). (See under "Parasitism.")
As is the case with every other vital function, so for the function
of metabolism we find a starting-point in the lowest and simplest
group of the protophyta, the chromacea. In their oldest forms, the
chroococcacea, the whole body is merely a blue-green, structureless,
globular plasma particle, growing by means of its plasmodomous power,
and splitting up as soon as it reaches a certain stage of growth.
There the miracle of life consists merely of the chemical process of
plasmodomism by photosynthesis. The sunlight enables the blue-green
phytoplasm to form new plasm of the same kind out of inorganic
compounds (water, carbonic acid, ammonia, and nitric acid). We may look
upon this process as a special kind of catalysis. In this case there
is absolutely nothing to be done by Reinke's "dominants," or conscious
and purposive vital forces. There are, as yet, no differentiated
physiological functions in these organisms without organs, and no
anatomically distinct members; and so their one vital activity, growth,
may very well be compared to the simple growth of inorganic crystals.
It has been pointed out repeatedly that the remarkable monera which
now play so important a part in biology as bacteria stand, in many
respects, quite apart from the ordinary vital phenomena of the
higher organisms. This is especially true of their metabolism, which
has the most striking peculiarities. Morphologically, many of the
bacteria cannot be distinguished from their nearest relatives and
direct ancestors, the chromacea, differing from them only in the
absence of coloring matter in the plasm. Many of them are simple,
globular, ellipsoid, or rod-shaped plasma particles, without any
visible organization or movement. Others move about by means of one or
more very fine lashes (like the flagellata). No real nucleus can be
discovered in the structureless plasma body. The very fine granules
which are found in some species, and the vacuole-formation that we see
in others, may be regarded as products of metabolism; and the same
may be said of the thin membrane or the thicker gelatinous envelope
which many of the bacteria secrete. This makes all the more remarkable
the peculiarity of their chemical constitution and the metabolism
determined thereby. The nitro-bacteria we have mentioned previously are
plasmodomous; the anaërobe bacteria (of butyric acid and tetanus) only
flourish where oxygen is excluded; the sulphur bacteria (_beggiatoa_)
secrete--by the oxydation of sulphuretted hydrogen--pure regulation
sulphur in the form of round granules. The ferruginous bacteria
(_leptothrix ochrocea_) store up oxyhydrate of iron (by the oxydation
of carbonic protoxide of iron). The saprogenetic bacteria cause
putrefaction, and the zymogenetic fermentation. Finally, we have the
very interesting pathogenetic bacteria which cause the most dangerous
diseases by the secretion of special poisons--toxins--festering,
small-pox, tetanus, diphtheria, typhus, tuberculosis, cholera, etc.
On account of their great practical importance, these bacteria have
of late been taken over by a special branch of biology, bacteriology.
But only a few of the many experts in this department have pointed out
the extreme theoretical significance which these zoomonera have for
the important questions of general biology. These structureless plasma
bodies show unmistakably that their vital activity is a purely chemical
phenomenon. Their great variety proves how manifold and complicated
must be the molecular composition of the plasm, even in these simplest
organisms.
The unicellular protophyta exhibit the same form of metabolism and
plasmodomism as the familiar green cells of the tissue-plants; but
in most of the protozoa we find special features of nutrition and
plasmophagy. The great class of the rhizopods is distinguished by the
fact that their naked plasma body can take in ready-formed solid food
at any point of its surface. On the other hand, most of the infusoria
have a definite mouth-opening in the outer wall of their unicellular
body, and sometimes a gullet-tube as well. Besides this cell-mouth
(_cytostoma_) we usually find also a second opening for the discharge
of indigestible matter, a cell-anus (_cytopyge_).
Metabolism in the tissue plants (metaphyta) forms a long gradation from
very simple to very complicated arrangements. The lowest and oldest
thallophyta, especially the simplest algæ, are not far removed from
the communities of protophyta, and, like these, are merely definitely
grouped colonies of cells. The social cells which form their most
rudimentary tissue are quite homogeneous, with no differentiation
beyond that of sex. The thallus or bed-formation consists in the
simplest specimens of plain or branched fine threads, consisting of
rows or chains of homogeneous cells (so _conferva_ among the green,
_ectocarpus_ among the brown, and _callithamnion_ among the red
algæ). Other algæ (such as the ulva) form thin leaf-shaped forms of
the thallus, a number of homogeneous cells lying side by side along a
level. In the larger algæ compact tissue-bodies are formed, in which
frequently firmer rows of cells exhibit the rudiments of fibres; and
the thallus divides, as in the cormophyta, into root, stalk, and
leaves. There is also a trophic differentiation, the fibres undertaking
special functions of nutrition (the conduction of the sap). The
same must be said of the mosses (_bryophyta_). Their lowest forms
(_ricciadinæ_) are close akin to the algæ; the highest mosses (the
_mnium_ and _polytrichum_, for instance) approach the cormophyta. Many
botanists comprise these lower plants--algæ, fungi, and mosses--under
the title of "cell-plants" (_cytophyta_), and oppose the higher
plants--ferns and flowering-plants--to them as "vascular plants"
(_angiophyta_), because they have complex fibres or sap vessels. This
distinction has a phylogenetic significance similar to the division
between cœlenteria and cœlomaria in the animal kingdom.
While most of the cell-plants either live in the water (algæ) or are
very simply organized on account of their saprophytic or parasitic
habits (fungi), the vascular plants mostly live on land, and have to
adapt themselves to much more complicated conditions. Their nutrition
is accordingly distributed among different functions, and special
organs have been evolved to discharge them. This is equally true of
the crytogam ferns (_pteridophyta_) and the phanerogam flowering
plants (_anthophyta_). The most important later acquisition which
distinguishes both groups from the lower cell-plants is the possession
of vascular or conducting fibres. These organs for conducting water
pass through the entire body of the vascular plant in the shape of
long tubes, formed by the combination of rows of cells; the cells
themselves die off, and their plasma content disappears. The stream of
water that rises constantly in these tubes is taken up by the roots,
conducted by the fibres to all parts, and given off (transpiration) by
the pores of the leaves. But these pores also serve for the breathing
of plants, being connected with the air-containing intercellular
passages; through these air-spaces, which serve for the aëration of
the higher plant-body, air and moisture can enter, and oxygen be given
off in respiration. Finally, many of the vascular plants have special
glands that serve for secretion (of oil, resin, etc.). In the higher
flowering plants this division of work among the various digestive
organs gives rise to a very complicated apparatus for nutrition. Among
the many remarkable structures that have been developed in this way by
adaptation to special conditions we may particularly note the organs
for catching and digesting insects in the insect-eating plants, the
European _drosera_ and _utricalaria_, and the tropical _nepenthas_ and
_dionæa_.
The long scale of evolutionary forms which we find in the tissue
animals (metazoa) leads up uninterruptedly from the simplest to
the most elaborate physiological functions and a corresponding
morphological complexity of organs. The two principal divisions of
the metazoa are chiefly distinguished by the circumstance that in the
cœlenteria one single system of organs, the gastro-canal system,
discharges the whole (or most part) of the partial functions of
nutrition; while in the cœlomaria they are usually distributed among
four different systems of organs, each of which is made up of a number
of organs. To an extent, we find once more in each great division
characteristic types of organization. However, comparative ontogeny
teaches us that all these various structures have been developed
from one simple fundamental form, as I have shown in my theory of the
gastræa (1872).
The older research into the origin of the nutritive apparatus in
the metazoa--especially its chief part, the alimentary or gastric
canal--had led to the erroneous belief that in several groups of the
metazoa it owed its origin to very different growth-processes, and
that particularly in the higher vertebrates (the amniotes) it was
a comparatively late product of evolution. On the other hand, the
comparative study of the embryology of the lower and higher animals
led me thirty-four years ago to the opposite conclusion, that a simple
gastric sac was the first and oldest organ of all the metazoa, and that
all the different forms of it had been developed from this primitive
type. I gave this view in my _Biology of the Sponges_ in 1872; and I
developed and established it in my _Studies of the Gastræa Theory_ in
1873. In the latter book I also worked out the important conclusions
that follow from this monistic reform of the theory of germinal layers
for the phylogenetic natural classification of the animal kingdom.
I began with the consideration of the simplest sponges (_olynthus_)
and cnidaria (_hydra_). The whole body of these lowest and oldest
of the cœlenteria is in essence nothing but a round, oval, or
cylindrical gastric vesicle, a digestive sac, the thin wall of which
consists of two simple layers of cells. The outer layer (the ectoderm
or skin-layer) is the covering layer of the external skin (epidermis);
it is the instrument of sensation and movement. The inner layer of
cells (entoderm or gastric layer) serves for nutrition; it clothes
the simple cavity of the sac, which admits the food by its opening
and digests it. This opening is the primitive mouth (_prostoma_ or
_blastoporus_), the inner cavity itself the primitive gut (_progaster_
or _archenteron_). I proved that there was the same composition in
the young embryos or larvæ of many of the lower animals, and showed
that the manifold and apparently very different embryonic form of all
the higher animals may be reduced to the same common type. To this I
gave the name of the "cup-embryo" or gastric larvæ (_gastrula_), and
concluded, in virtue of the biogenetic law, that it is the palingenetic
reproduction of a corresponding ancestral form (the _gastræa_)
maintained until the present by heredity. It was not until much later
(1895) that Monticelli discovered a modern gastræad (_pemmatodiscus_)
which corresponds completely to this hypothetical ancestor (see the
last edition of my _Anthropogeny_, fig. 287). The simplest living forms
of the sponges (_olynthus_) and the cnidaria (_hydra_) only differ from
this hypothetical primitive form of the gastræa by a few secondary and
subsequently acquired features.
The classes of the lower animals which we comprise under the name
cœlenteria (or cœlenterata in the widest sense) generally agree
in having all the functions of nutrition accomplished exclusively (or
for the most part) by a single system of organs, the gastro-canal or
gastro-vascular system. From their common stem-group, the gastræads,
three different stems have been evolved--the sponges, cnidaria, and
platodes. All these cœlenteria have three features in common: (1)
The gastric canal or tube has only one opening--the primitive mouth,
which serves at once for admitting food and ejecting indigestible
matter; there is no anus; (2) there is no special body-cavity
(_cœloma_) distinct from the gastric tube; (3) there is also no
trace of a vascular system. All cavities that are found in these lower
animals besides the digestive gut-cavity are direct processes from it
(with the exception of the nephridia in the platodes).
While the simple digestive gut is the sole organ of nutrition in the
stem-group of the gastræads, we find other structures co-operating in
the rest of the cœlenteria. The characteristic stem of the sponges
is distinguished by the piercing of the wall of the gastric vesicle
with several holes. Through these water pours into the body, bringing
with it the small particles of food which are received and digested
by the ciliated cells of the entoderm; the water emerges again by the
mouth-opening (_osculum_). The best-known of the sponges is the common
bath-sponge (_euspongia officinalis_), the horny skeleton of which
we use daily in washing. In these and most other sponges the large,
unshapely body is traversed by a number of branching canals, on which
there are thousands of tiny vesicles, produced by the multiplication
of a simple gastric vesicle of the primitive sponge (_olynthus_). Each
of these ciliated chambers is really a tiny gastræa, a "person" of the
simplest character (_cf._ chapter vii.). Hence we may regard the whole
sponge-body as a gastræad-stock (_cormus_).
The large group of the cnidaria offers a long series of evolutionary
stages, from very small and simple to very large and elaborate forms.
Some of them remain at a very low stage, as does our common green
fresh-water polyp (_hydra viridis_), which only differs from the
gastræa by a few variations in tissue and the formation of a crown of
feelers about the mouth. Most of the polyps form stocks (_cormi_),
the individuals shooting out buds which remain joined to the mother
animal. In these and all the other stock-forming animals the nutrition
is communistic; all the food that the individuals get and digest is
conducted by tubes to the common fund and equally distributed. In all
the larger cnidaria the body-wall becomes thicker, and is traversed by
branching gastro-canals; these convey the nutritive fluid to all parts
of the body.
While the fundamental type in the cnidaria is radial (determined
by the crown of radiating feelers or tentacles that surrounds the
mouth), it is bilateral-symmetrical in the platodes or "flat-worms"
(_plathelminthes_). In this animal-stem, moreover, the lowest forms,
the platodaria (also called _cryptocœla_ and _acæla_) come very
close to the gastræa. But most of the platodes are distinguished from
the rest of the cœlenteria by the formation of a pair of nephridia
(renal canals or water-vessels), thin tubes which, as excretory organs,
remove from the body the unusable products of metabolism, the urine.
Here we have a second organ of nutrition, the gut tube, added to the
first. In the lower platodes this remains very simple. As a rule, a
gullet tube (pharynx) is formed by the hollowing out of the mouth, as
in the corals; and as in the case of the latter branched canals, which
conduct the nutritive sap from the stomach to distant parts of the
body, grow out of the stomach, in the larger coil-worms (_turbellaria_)
and suction-worms (_trematodes_). On the other hand, the gut atrophies
in the tape-worms (_cestodes_); as these parasites live in the
intestines or other organs of animals, they can obtain their nutritive
sap directly from them through the surface of the skin.
The more highly organized cœlomaria differ from the simpler
cœlenteria chiefly by the greater complexity in the structure
and functions of their apparatus of nutrition. As a rule, these
functions are divided between four groups of organs, which are not yet
differentiated in the cœlenteria--namely: 1, organs of digestion
(gastric system); 2, organs of circulation (vascular system); 3, organs
of breathing (respiratory system); and 4, organs of excretion (renal
system). Moreover, in the cœlomaria the gastric canal has usually
two openings, the mouth and the anus. Finally, they all have a special
body-cavity (_cœloma_); this is quite separate from the gastric
canal, which is suspended in it, and serves for the formation of the
sexual cells. It is formed in the embryo by the hollowing out and
cutting off of a pair of sacs (cœlom-pouches) from the gut near the
mouth; the pouches touch, and then coalesce, as their division-walls
break down. If a part of the dividing wall remains, it serves as
mesentery to fasten the gut to the body-wall. The action of the four
groups of alimentary organs remains very simple in the lowest and
oldest cœlomaria, the worms (_vermalia_); but in the other higher
animals, which have been evolved from these, they have very varied and
often complicated features.
In the great majority of the cœlomaria the gastric system forms a
highly differentiated apparatus, composed, as in man, of a number of
different organs. The food is usually taken in by the mouth, ground
up by the jaws or the teeth, and softened with saliva, which the
salivary glands pour into the cavity of the mouth. From the mouth
the pulpy food passes in swallowing into the gullet, which often has
glandular appendages, and from this through the narrow esophagus into
the stomach. This most important part of the alimentary apparatus is
often divided into several sections, one of which (the masticating
stomach) is armed with teeth and prepared for a further triturition
of solid pieces, while the other (the glandular stomach) produces the
dissolving gastric juice. The liquefied food (_chylus_) then passes
into the small intestine (_ileum_), which has to absorb it, and is as a
rule the longest section of the alimentary canal. A number of different
digestive glands open into this intestine, the most important of them
being the liver. The small intestine is often sharply distinguished
from the large intestine (_colon_), the last large section of the
alimentary canal; into this also a number of glands and blind
intestines open. The last portion of it is called the _rectum_, and
this removes the indigestible remnants of the food (_fæces_) through
the anus.
This general plan of the alimentary system, which is common to most
of the cœlomaria in its chief features, is very much modified in
the various groups of these animals and adapted to their several
conditions of nutrition. The simplest structures are found in many of
the vermalia; the lowest forms of these, the rotifers, and especially
the gastrotricha, still closely resemble their platode ancestors, the
turbellaria. The higher type of animal-stems which have been evolved
from them are partly distinguished by special structures. Thus the
mollusks have a characteristic masticating apparatus; on their tongue
there is a hard plate (_radula_) armed with a number of teeth, which
grinds against a hard upper jaw, and so breaks up the food. In most of
the articulates this work is done by side-jaws, which consist of hard
rods and represent modified bones. The vertebrates and the closely
related tunicates are distinguished by the conversion of the first
sections of the alimentary canal into a characteristic respiratory
apparatus (gills). But the construction of the various sections of
the gastro-canal also varies a good deal in the small groups of
the cœlomaria, as it depends to a great extent on the nature of
the food and the conditions in which it is got and prepared. The
largest expenditure of mechanical and chemical energy is needed for
a voluminous solid vegetal diet. Hence the alimentary canal and its
many appendages are longest and most complicated in the plant-eating
snails, leaf-eating insects, and grass-eating ruminants. On the other
hand, they are shortest and simplest in parasitic cœlomaria, which
derive their fluid food already prepared from the contents of another
animal's intestines. In these cases the gut may altogether atrophy; as
in the _acanthocephala_ among the vermalia, the _entoconcha_ among the
mollusks, and the _sacculina_ among the crustacea.
The greater the extent of the body, and the more complex the
organization of the higher animals, the more necessary it is to have
an orderly and regular distribution of the nutritive fluid to all
parts. In the cœlenteria this work is accomplished by the gastric
canals (side branches from the gut, opening into its cavity) but in
the cœlomaria it is done much better by means of blood-vessels
(_vasa sanguifera_). These canals do not communicate directly with the
gastro-canal, but are formed independently of it in the surrounding
parenchyma of the mesoderm. They take up the filtered and chemically
improved food-fluid, which transudes through the intestinal walls, and
conduct it in the form of blood to all parts of the body. This blood
generally contains millions of cells, which are of great importance
in metabolism. The blood-cells of the lower cœlomaria are usually
colorless (leucocytes), while those of the vertebrates are mostly red
(rhodocytes).
The circulation of the blood in most of the cœlomaria is effected
by a heart, a contractile tube, formed by the local thickening of
a skin-vessel, which contracts and beats regularly by means of its
muscular bands. Originally two of these skin-vessels were developed in
the abdominal wall--a dorsal vessel in the upper and ventral vessel
in the lower wall (as in many of the vermalia). The heart is formed
from the dorsal vessel in the mollusks and articulates, but from the
ventral in the tunicates and vertebrates. The arteries are the vessels
which conduct the blood from the heart; those which conduct it from the
body to the heart are the veins. The finest branchlets of both kinds
of vessels, which form the connecting link between them, are called
capillaries; these immediately effect the interchange of matter in the
tissues by osmosis. The blood-vessels co-operate very closely with the
respiratory organs.
The interchange of gases in the organism, which we call breathing or
respiration--the taking in of oxygen and giving out of carbonic-acid
gas--does not require special organs in the lower animals. In these
it is accomplished by epithelial cells, which clothe the surface of
the body--the ectoderm of the outer skin layer and the entoderm of
the inner gut-covering. As nearly all these cœlenteria live in the
water, or (as parasites) in some fluid that contains air, and as these
fluids are constantly pouring in and out of the body, the exchange of
gases is accomplished at the same time. But in the higher animals this
is rarely found, only in the small animals of simple construction (such
as the rotifers and other vermalia, and the smallest specimens of the
mollusca and articulata). The majority of these cœlomaria attain a
considerable size, and so require special organs; these afford a larger
surface for the exchange of gases in the limited space, and accomplish
a very peculiar chemical work as the localized organs of respiration.
They fall into two groups according to the nature of the environment;
gills for breathing in water and lungs for breathing on land. The
latter take the oxygen directly from the atmosphere, and the former
from the water, in which atmosphere air is contained in solution.
The instruments of water-respiration which we call gills (_branchiæ_)
are generally attenuated parts or processes of the outer skin or the
inner gastric skin; hence we distinguish the two chief forms, external
and internal gills. Both are richly provided with blood-vessels which
bring the blood from the body for the purpose of aëration. Cutaneous
or external gills are especially found in the vertebrates, in the form
of threads, combs, leaves, pencils, tufts of feathers, etc., which are
drawn out from the entoderm as local processes of the outer skin, and
afford a wide surface for the interchange of gases between the body
and the water. In the mollusca there are usually a pair of comb-shaped
gills near the heart; in the articulates there are several pairs,
repeated in the different segments of the body. Gastric or internal
gills are peculiar to the vertebrates and the next-related tunicates,
with a small group of the vermalia, the enteropneusta. In these the
fore-gut or head-gut is converted into a gill-organ, the wall of which
is pierced with gill-fissures; the water that has been taken in by the
mouth passes away through the outer openings of these fissures. In the
lower aquatic vertebrates (acrania, cyclostoma, and fishes) the gills
are the sole organs of breathing; in the higher animals, that live in
the air, they fall into disuse, and their place is taken by lungs.
Nevertheless, heredity is so tenacious that we find from three to five
pairs of rudimentary gill-clefts in the embryo right up to man, though
they have long since ceased to have any function. This is one of the
most interesting of the palingenetic facts that prove the descent of
the amniotes (including man) from the fishes.
The group of the aquatic echinoderms has some very peculiar features of
respiration. Their body possesses an extensive water-duct, which takes
in the sea-water and returns it by special openings (skin-pores or
madreporites). The many branches of these water-vessels or ambulacral
vessels fill with water, especially the tiny feelers or feet, which
extend from the skin in thousands; they serve at once for movement,
feeling, and breathing. But many of the echinoderms have also special
gills--the star-fish have small finger-shaped cutaneous gills on
the back, the sea-urchins special leaf-shaped ambulacral gills, the
sea-cucumbers internal gastric gills (tree-shaped branching internal
folds of the rectum).
The organs of air-breathing are called, in general, lungs (_pulmones_).
Like the organs of water-breathing, they are formed sometimes from
the external and sometimes from the internal covering of the body.
Cutaneous or external lungs are found in several groups of the
vertebrates. Among the mollusks the land-dwelling lung-snails have
acquired a lung-sac by change in the work of the gill cavity: among
the articulata the lung-spiders and scorpions have two or more
trachea-lungs; that is to say, cutaneous sacs, in which are enclosed
fanwise a number of trachea-leaves. In the other air-breathing
articulates (tracheata) we find, instead of these simple or branched,
and often bushlike, air-tubes (_tracheæ_), which spread through the
whole body and conduct the air direct to the tissues. They take the
air from without by special air-holes in the skin (_stigmata_ and
_spiracula_). The myriapods and insects generally have numbers of
air-holes; the spiders only one or two, more rarely four, pairs. When
these air-tube animals return to an aquatic life (as happens with the
larvæ of various groups of insects), the outer air-holes close up, and
new thread-shaped or leaf-shaped trachea-gills are formed, which take
the air from the surrounding water by osmosis. The oldest and lowest
tracheata are the primitive air-tube animals, or protracheata, and form
the link between the older annelids and the myriapods. They have a
number of clusters of short air-tubes distributed over the whole skin,
and it is clear that these have been evolved from simple skin-glands by
change of function.
Gastric or internal lungs are only found in the higher animals, to
which we give the name of quadrupeds (or _tetrapoda_), the amphibia and
amniotes, and their fishlike ancestors, the dipneusta. These internal
lungs are sac-shaped folds of the fore-gut, formed originally from the
swimming-bladder (_nectocystis_) of the fishes by change of function.
This air-filled bladder, a sac-shaped appendage of the gullet, merely
serves the purpose of a hydrostatic organ, by varying the specific
weight, in the fishes. When the fish wishes to descend it contracts
the bladder and becomes heavier; it rises to the top by inflating it
again. The lungs were formed by the adaptation of the blood-vessels
in the wall of the swimming-bladder to the interchange of gases. In
the oldest living lung-fishes (_ceratodus_) it is still a simple sac
(_monopneumones_=one-lunged); in the others the simple gullet-cavity
divides early into a pair of sacs (_dipneumones_, two-lunged). The
wind-pipe (_trachea_--not to be confused with the organ of the same
name in the tracheata) is formed by the lengthening of their stalk
and strengthening of it with cartilaginous rings. At the anterior end
of the trachea we find already formed in the amphibia the larynx, the
important organ of voice and speech.
The function of removing unusable matter is not less important to the
organism than breathing. Just as breathing gets rid of the poisonous
carbonic acid, so the kidneys remove fluid and solid excreta in the
shape of urine; these are partly acid (uric acid, hippuric acid, etc.),
partly alkaline (urea, guanine, etc.). In most of the cœlomaria
special organs for removing these would be superfluous, as this is
accomplished (like breathing) by the stream of water that is constantly
passing through the whole body. But with the platodes we begin to
find important excretory organs in the nephridia, a pair of simple
and ramified canals which lie on either side of the gut, and open
outward. These primitive renal canals are transmitted by the platodes
to the vermalia, and by these to the higher stems of the cœlomaria.
In the latter they generally open by special funnels into the inner
body-cavity, which serves as first receptacle for the urine. Their
outer opening sometimes (primarily) goes through the outer skin at the
back (excretory pores), sometimes (secondarily) to the rectum, and
so out through the anus. The oldest articulates, the annelids, have
a pair of nephridia in each segment of the body; each renal canal,
or segmental canal, consists of three sections, an inner funnel
which opens into the body-cavity, a middle glandular section, and an
external bladder that ejects the urine by contraction. The disposition
of the renal system in the internally articulated vertebrates is very
similar to this; but now complicated structures begin to appear, a
pair of compact kidneys (_renes_), which are made up of a number of
branching nephridia. Three generations of kidneys succeed each other,
as phylogenetic stages of evolution--first the primary fore-kidneys
(_protonephros_), in the middle the secondary primitive kidneys
(_mesonephros_), and last the tertiary after-kidneys (_metanephros_).
The latter are only reached in the three highest classes of
vertebrates, reptiles, birds, and mammals. Mollusks also have a couple
of compact kidneys. They are developed from a pair of nephridia, the
funnels of which open internally into the heart-pouch (the remainder of
the reduced body-cavity); at the back they open outward. The crustacea
also have generally a pair of renal canals. On the other hand, the
protracheata (the stem-forms of the air-tube animals) have segmental
nephridia, a pair to each joint inherited from their annelid ancestors.
The rest of the tracheata, the myriapods, spiders, and insects, have,
instead of these, Malpighi tubes, funnel-shaped glands that arise from
the entodermal rectum, sometimes one pair or less, sometimes a number
in a cluster.
While most plants are purely plasmodomous, and most animals
plasmophagous, there are nevertheless in both organic kingdoms a
number of species (especially the lower) whose metabolism has assumed
peculiar forms by their relations to other organisms. To this class
belong especially the saprosites and parasites. By saprosites are
understood those plants and animals which feed entirely or mostly
on the corpses of other animals, or the decomposed matter which is
unfit for the food of higher animals. Among the unicellular protists
many of the bacteria, especially, belong to this class, and also many
fungilla (_phycomycetes_); among the metaphyta the fungi (mycetes),
and among the metazoa the sponges. I have already spoken of the many
peculiarities of metabolism in the ubiquitous bacteria; while many of
them cause putrefaction, they at the same time feed on the parts of
other organisms which have died. The fungi feed for the most part on
the decayed remains of plants and the products of putrefaction which
accumulate on the ground. In this character of scavengers they play the
same important part on land as the sponges do at the bottom of the sea.
But a number of small groups of the higher plants and animals have, as
a secondary habit, turned to saprositism. Among the metaphyta we have
especially the monotropea (to which our native asparagus, _monotropa
hypopitys_, belongs) and many orchids (_neottia_, _corallorhiza_). As
they find their plasm directly in the decayed matter in the woods,
they have lost their chlorophyll and green leaves. Among the metazoa
many of the vermalia, and some of the higher animals, such as the
rain-worm and many tube-dwelling annelids (the mud-eaters, _limicolæ_),
etc., live on putrid matter. The organs which their nearest relatives
use for obtaining, breaking up, and digesting food (eyes, jaws,
teeth, digestive glands) have been entirely or mostly lost by these
saprosites. Many of them form a transitional type to the parasites.
By parasites, in the narrower sense, we understand, in modern biology,
only those organisms which live on others and derive their nourishment
from them. They are numerous in all the chief divisions of the plant
and animal kingdoms, and their modifications are of great interest
in connection with evolution. No other circumstance has so profound
an influence on the organism as adaptation to a parasitic existence.
Moreover, there is no other section in which we can follow, step
by step, the course of the degeneration which is caused, and show
clearly the mechanical nature of the process. Hence the science of
parasites--parasitology--is one of the soundest supports of the theory
of descent, and provides an abundance of the most striking proofs of
the much-contested inheritance of acquired characteristics.
Among the unicellular organisms, the bacteria are the most conspicuous
instances of manifold adaptation to parasitic habits. As we count
these unnucleated protozoa among the oldest and simplest organisms,
and trace them directly by metasitism to the plasmodomous chromacea,
it is very probable that they turned to parasitism very early in the
history of life. Even a part of the monera (in which group we must
place the bacteria on account of their lack of a nucleus) found it
convenient and advantageous to prey on other protists and assimilate
their plasm directly, instead of going through the laborious process
of carbon assimilation themselves in the hereditary fashion. This is
also true of the large class of the sporozoa or fungilla (_gregarinæ_,
_coccidia_, etc.), real nucleated cells, which have adapted themselves
in various ways to parasitic habits. Many of them live in the
rectum, the cœlum, or other organs of the higher animals (the
gregarinæ, especially in the articulates); others in the tissues (for
instance, the sarcosporidia in the muscles of mammals, the coccidia
and myxosporidia in the liver of vertebrates). A good many of them
are "cell-parasites," and live inside the cells of other animals,
which they destroy; such are the hœmosporidia, which destroy the
blood-cells in man, and so cause intermittent fever.
Among the multicellular metaphyta it is particularly the fungi that
have taken to parasitism in various ways. Many of them are, as is
known, the most dangerous enemies of the higher animals and plants. The
various species of fungi cause certain diseases by their poisonous
(chemical) action on the tissues of their host. It is well known how
our most important cultivated plants, the vine, potato, corn, coffee,
etc., are threatened by fungoid diseases; and this is also true of many
of the lower and higher animals. It is probable that the fungi have
been evolved polyphyletically by metasitism from the algæ.
Among the higher metaphyta we find parasitism in many different
families, especially orchids, rhinanthacea (_orobranche_,
_lathraca_), convolvulacea (_cuscuta_), aristolochiacea, loranthacea
(_viscum_, _loranthus_), rafflesiacea, etc. These various kinds of
flowering-plants often grow to resemble each other by convergence
(that is to say, by their common adaptation to parasitic life); they
lose their green leaves, the plasmodomous chlorophyll of which is of
no further use to them. Frequently rudimentary leaves are left on them
in the form of colorless scales. For the purpose of clinging to the
plants they live on, and penetrating into their tissues, they evolve
special clinging apparatus (haustoria, suctorial cups, creepers). Their
stalks and roots are also modified in a characteristic way. The whole
productive force of these parasites is expended on their sexual organs;
_rafflesia_ has the largest flowers there are, more than a yard in
diameter.
Parasitism in the metazoa (in all groups) is even more frequent and
interesting than in the metaphyta. The mollusks and echinoderms
show the least disposition for it, and the platodes, vermalia, and
articulates the most. Even among the gastræada, the common ancestral
group of the metaphyta, we find parasites (kyemaria and gastremaria).
The protection they find inside their hosts is probably the reason
why these oldest of the metazoa have remained unchanged to the
present day. Real parasites are not numerous among the sponges and
cnidaria. But they are very numerous among the platodes. The suctorial
worms (_trematodes_) live partly externally (as ectoparasites)
on other animals and partly inside them (as endoparasites), and
produce serious diseases in them. They have lost the vibratory
coat of their free-living ancestors, the turbellaria, and acquired
clinging apparatus instead. The tape-worms (_cestodes_), which live
entirely in the interior of other animals, and are descended from the
suctorial worms, have lost their gastro-canal; they are nourished by
imbibition through the skin. The same degeneration is found in the
itchworms (_acanthocephala_) among the vermalia, the parasitic snails
(_entoconcha_) among the mollusks, and the root-crabs (_rhizocephala_)
among the crustacea.
The class of crustacea affords the most numerous and most instructive
examples of degeneration through parasitism, because in this class it
is found polyphyletically in very different orders and families, and
because their highly organized body shows every stage of degeneration
together in the different organs. The free-living crustacea generally
move about very rapidly and ingeniously; their numerous bones are
well jointed and excellently adapted for the most varied methods of
locomotion (running, swimming, climbing, digging, etc.); their organs
of sense are highly developed. As these are no longer used when
they take to parasitism, they atrophy and gradually disappear. The
younger crustacea all proceed from the same characteristic form of
the _nauplius_, and swim freely about; later, when they settle down
to parasitic habits, their organs of sense and locomotion atrophy. As
Fritz Müller-Desterro showed in his famous little work, _For Darwin_
(1864), forty years ago, the crustacea afford most luminous proofs of
the theory of descent and selection, and of progressive heredity and
the biogenetic law. These facts are the more important as the crab
undergoes the same degeneration by parasitic habits in a number of
different orders and families.
From parasitism we must entirely distinguish that intimate life-union
of two different organisms which we called symbiosis or mutualism.
Here we have an association of two living things for their mutual
benefit, while the parasite lives entirely at the expense of his
host. Symbiosis is found among the protista, being very wide-spread
among the radiolaria. In the gelatinous envelope (_calymma_) which
encloses the central capsule of their unicellular bodies we find a
number of motionless yellow cells (_zooxanthella_) scattered. These
are protophyta or (as it is said) "unicellular algæ" of the class of
paulotomea (_palmellacea_). They receive protection and a home from the
radiolaria, grow plasmodomously, and multiply by rapid segmentation.
A large part of the starch-flour and the plasm which they form by
carbon-assimilation goes as food directly to the radiolarium-host; the
other part of the xanthella goes on growing and multiplying. Similar
yellow zooxanthella or green zoochlorella are found as symbionta in the
tissues of many animals. Our common fresh-water polyp (_hydra viridis_)
owes its green color to the zoochlorella which live in great numbers
on the ciliated cells of its entoderm (the digestive gut-epithelium).
In general, however symbiosis is rarer in the metazoa than in the
metaphyta. In the latter case it is the fundamental feature of a whole
class of plants, the lichens. Each lichen consists of a plasmodomous
plant (sometimes a protophyte, sometimes an alga) and a plasmophagous
fungus. The latter affords a home, protection, and water to the green
alga, which repays the service by providing food.
XI
REPRODUCTION
Reproduction and generation--Sexual
and asexual reproduction--Superfluous
growth--Monogony--Self-cleavage--Budding--Formation of
spores--Amphigony--Ovum and sperm-cell--Hermaphrodite formation
and separation of the sexes--Hermaphrodism and gonochorism
of the cells--Monoclinism and diclinism--Monœcism and
diœcism--Alternation of sex-division--Sexual glands of
the histona--Hermaphroditic glands--Sexual ducts--Generative
organs--Parthenogenesis--Pædogenesis--Metagenesis--Heterogenesis--
Strophogenesis--Hypogenesis--Hybridism--Generation of hybrids and
the species--Graduation of forms of reproduction.
While nutrition secures the maintenance of the organic individual,
reproduction insures that of the organic species, or the group of
definite forms which we distinguish from others by the name "species."
All individuals are more or less restricted in the duration of their
lives, and die off after the lapse of a certain time. The succession
of individuals, connected by reproduction and belonging to a species,
makes it possible for the specific form itself to last for ages. In
the end, however, the species is temporary; it has no "eternal life."
After existing for a certain period, it either dies or is converted by
modification into other forms.
The rise of new individuals by reproduction from parent organisms
is a natural phenomenon with definite time-restriction. It cannot
have continued from eternity on our planet, as the earth itself is
not eternal, and even long after its formation was incapable of
supporting organic life on its surface. This only became possible when
the surface of the glowing planet had sufficiently cooled for liquid
water to settle on it. Until this stage carbon could not enter into
those combinations with other elements (oxygen, hydrogen, nitrogen, and
sulphur) which led to the formation of plasm. As I intend to deal with
this process of _archigony_, or spontaneous generation, in a special
chapter, I leave it for the present, and confine myself to the study of
_tocogony_, or parental generation.
* * * * *
The various forms of tocogony, or the reproduction of living things,
are generally divided into two large groups; on the one hand there is
the simple form of asexual generation (monogony), and on the other the
complex form of sexual generation (amphigony). In asexual generation
the action of one individual only is needed, this providing a product
of transgressive (redundant) growth which develops into a new organism.
In sexual generation it is necessary for two different individuals to
unite in order to produce a new being from themselves. This amphigony
(or _generatio digenea_) is the sole form of reproduction in man and
most of the higher animals. But in many of the lower animals and most
of the plants we find also asexual multiplication, or monogony, by
cleavage or budding. In the lowest organisms, the monera and many of
the protists, fungi, etc., the latter is the only form of propagation.
Strictly speaking, monogony is a universal life-process; even the
ordinary cell-cleavage, on which depends the growth of the histona, is
a cellular monogony. Hence historical biology must say that monogony
is the older and more primitive form of parental generation, and
that amphigony was secondarily developed from it. It is important to
emphasize this because, not only some of the older writers, but even
some recent ones, regard sexual generation as a universal function of
organisms, and declare that it dates from the very beginning of organic
life.
The complex and frequently very intricate phenomena of sexual
generation, as we find them in the higher organisms, become
intelligible to us when we compare them with the simpler forms of
asexual generation at the lowest stages of life. We then learn that
they are by no means unintelligible and supernatural marvels, but
natural physiological processes, which, like all others, may be traced
to the action of simple physical forces. The form of energy which
lies at the root of all tocogony is _growth_ (_crescentia_). And as
this phenomenon is also the cause, in the form of gravitation, of the
formation of crystals and other inorganic individuals, we do away with
another of the boundaries which people would establish between organic
and inorganic nature. Reproduction is a kind of nutrition and growth of
the organism beyond the individual standard, building up a part of it
into a whole. This _limit_ of individual size is determined for each
species by two factors--the inner constitution of the plasm, which is
inherited, and the dependence on the outer environment, which controls
adaptation. When this limit has been passed, the transgressive growth
takes the form of reproduction. Every species of crystal has also a
definite limit of growth; when this is passed, new crystal-individuals
are formed in the mother-water on the old individual, which grows no
further.
Asexual or monogenetic tocogony (also called "vegetative
multiplication") is always effected by a single organic individual,
and so must be traced to its transgressive growth. When this affects
the entire body as a total growth, the whole dividing into two or
more equal parts, we call the monogenetic process division (or
segmentation). But when the growth is partial, and affects only a
part of the individual, or when this special part separates from
the generating organism in the form of a bud (_gemma_), the process
is called budding or gemmation (_gemmatio_). Hence the essential
difference between the two forms of generation is that in division the
parent disappears in its partial products (children); these are of
the same age and form. But in budding the generating parent retains
its individuality; it is larger and older than the young bud. This
important difference between division and gemmation, which is often
overlooked, holds good both for protists (unicellulars) and histona
(multicellulars). The fact that in division the individual as such
is destroyed is a sufficient refutation of Weismann's theory of the
immortality of the unicellulars. (See above, and also the _Riddle_,
chapter xi.)
Reproduction by division is by far the most common of all forms of
propagation. It is the normal form of monogony, not only in many of the
protists, but also in the tissue-cells which compose the tissues of
the histona. It is, moreover, the sole method of propagation for most
of the monera, both chromacea and bacteria, which are in consequence
often comprised under the title of "cleavage-plants" (_schizophyta_).
Self-cleavage is also found among the higher multicellular
organisms--namely, the cnidaria (polyps, medusæ). It usually takes the
form of division into two parts (_dimidiatio_ or hemitomy), the body
splitting into two equal halves. The plane of division is sometimes
indefinite (fragmentary-cleavage), sometimes coincident with the long
axis (length-cleavage), sometimes with the transverse axis, vertical
to the long axis (transverse-cleavage), and less frequently with a
diagonal axis (oblique-cleavage). When the segmentation of a cell
proceeds so rapidly that the transverse-cleavage follows immediately
on the length-cleavage, and the two are at length made to coincide,
twofold division is changed into fourfold division. And when the
process is repeated in quick succession, and the body falls at last
into a number of small and equal pieces, we have manifold division
(polytomy); as in the spore-formation of the sporozoa and rhizopoda,
and in the embryonic sac of the phanerogams.
Asexual propagation by budding is chiefly distinguished from
segmentation by the fact that the determining transgressive growth is
only partial in the one and total in the other. The bud produced is,
therefore, younger and smaller than the parent from which it issues;
the latter may replace the lost part by regeneration and produce a
number of buds simultaneously or successively without losing its
individuality (whereas this is destroyed in division). Propagation
by budding is rare among the protists, and more common among the
histona--that is, with most of the tissue-plants and the lower,
stock-forming, tissue-animals (cœlenteria and vermalia). Most stocks
(cormi) are formed by a sprout or person shooting out buds which remain
united to it. The layer and shoots of tissue-plants are detached
buds. The two chief kinds of gemmation are terminal and lateral.
Terminal budding takes place at the end of the long axis, and is not
far removed from transverse division (for instance, the strobilation
of the acraspedæ medusæ and the chain tape-worms). Lateral budding is
much more common; it determines the branching of trees and generally of
complex plants, and also of the tree-shaped stocks of sponges, cnidaria
(polyps, corals), bryozoa, etc.
A third form of asexual reproduction is the formation of spores or
"germ-cells," which are usually produced in great numbers inside the
organism, then detached from it, and developed into new organisms
without needing fertilization. The spores are sometimes motionless
(rest-spores or paulospores); sometimes they have one or more lashes
which enable them to swim about (rambling-spores or planospores).
This monogenetic propagation is very common among the protists, both
protophyta and protozoa. Among the latter the sporozoa (gregarinæ,
coccidia, etc.) are remarkable for the passing away of the whole
unicellular organism in the formation of spores; in this case and
in many of the rhizopods (_mycetozoa_) the process coincides with
manifold cell-division. In other cases (radiolaria, thalamophora)
only a portion of the parental cells is used for the production of
spores. Spore-formation is very common among the cryptogams; here it
usually alternates with sexual propagation. The spores are generally
formed in special spore-capsules (sporangia). In the flowering plants
(anthophyta) sporogony has disappeared. It is found at times in the
tissue-animals (in the fresh-water sponges); in this case the sporangia
are called _gemmulæ_.
The essential feature of sexual generation is the coalescence of two
different cells, a female ovum (egg-cell) and a male sperm-cell.
The simple new cell which arises from the blending of these is the
stem-cell (_cytula_), the stem-mother of all the cells that make up
the tissues of the histon. But even among the unicellular protists
we find in many places the beginnings of sexual differentiation; it
is foreshadowed in the blending or copulation of two homogeneous
cells, the gameta. We may conceive this process, or zygosis, as a
peculiar and very favorable kind of growth, that is connected with a
rejuvenescence of the plasm; the latter is enabled to propagate by
repeated cleavage through the mixing of the two different plasma-bodies
on either side (_amphimixis_). When these two gameta become unequal
and differ in size and shape, the larger female body is called the
macrogameton or macrogonidion, and the smaller, male part, the
microgameton or microgonidion. Among the histona the first is called
the egg-cell (_ovulum_), and the latter the sperm-cell (_spermium_, or
_spermatozoon_). As a rule the latter is a very mobile ciliated cell,
the former an inert or amœboid cell. The vibratory movements of the
sperm-cells serve for approaching the ovulum in order to fertilize it.
The qualitative difference between the two copulating sexual cells
(_gonocyta_), or the chemical difference between the ovoplasm of the
female and the sperm-plasm of the male cell, is the first (and often
the only) condition of amphigony; subsequently we find in addition (in
the higher histona) a very elaborate apparatus of secondary structures.
With this chemical difference is associated a peculiar double form of
sensitive perception and an attraction based thereon, which is called
sexual chemotaxis or erotic chemotropism. This "sex-sense" of the two
gonocyta, or elective affinity of the male androplasm and the female
gynoplasm, is the cause of mutual attraction and union. It is very
probable that this sexual sense-function, akin to smell or taste, and
the movements it stimulates, are located in the cytoplasm of the two
sex-cells, while heredity is the function of the caryoplasm of the
nucleus. (_Cf._ the _Anthropogeny_, chapters vi. and vii.)
The sexual difference between the two forms of gonoplasm, the ovoplasm
of the female and spermoplasm of the male cell, is noticeable at the
very beginning of sexual differentiation in the different sizes of
the copulating gameta, and later in their increasing divergence as to
shape, composition, movement, etc. It leads further to the distribution
of the germinal regions (in which the sex-cells are formed) into two
different individuals. When the ovum and the sperm-cell are produced in
one and the same individual, we call this an hermaphrodite; and when
they are formed in two different individuals (male and female), we
call them monosexual, or gonochorists. In accordance with the various
stages of individuality which we distinguished above (chapter vii.), we
may indicate the following stages of hermaphrodism and gonochorism.
Some groups of protists, especially the highly organized ciliated
infusoria (_ciliata_), are distinguished by having a separation of
male and female plasm within the unicellular organism. The ciliata
propagate, as a rule, in large numbers by repeated division (by
indirect cell-cleavage). But this monogony has its limits, and has to
be interrupted from time to time by amphigony, a rejuvenation of the
plasm, which is effected by the conjugation of two different cells and
the partial destruction of their nuclear matter. By conjugation is
meant the partial and momentary union of two different unicellulars,
while copulation is a total and permanent coalescence. When two
ciliated infusoria conjugate they place themselves side by side, and
connect for a time by means of a bridge of plasm. A part of the nucleus
of each has already divided into two portions, one of which functions
as the female standing-nucleus (_paulocaryon_) and the other as the
male travelling-nucleus (_planocaryon_). The two mobile nuclei enter
the plasm-bridge, and move through it, pushing against each other,
into the body of the opposite cell; they then coalesce with the deeper
lying standing-nucleus. When a fresh nucleus has been thus formed (by
_amphimixis_) in each of the copulating cells, they again separate. The
two rejuvenated cells have once more acquired the power to propagate
for a long time by division.
This peculiar hermaphroditic formation of the cells, which
distinguishes the ciliated infusoria and some other protists, and
which we now know in its smallest details through the investigations
of Richard Hertwig, Maupas, and others, is especially interesting
because it proves that the chemical difference between the female
gynoplasm and the male androplasm can be found within a single cell.
This erotic division of labor is so important that formerly it was
universally ascribed to two different cells. Recent accurate research,
penetrating into the smallest visible processes of fertilization, has
shown that the essential feature in the formation of a fresh individual
(the stem-cell) is the blending of equal portions (hereditary parts)
of the male and female nuclei; the caryoplasm of the two copulating
cells is the vehicle of heredity from the parents. The cytoplasm of the
cell-body, on the other hand, serves the purposes of adaptation and
nutrition. As a rule the cell-body of the ovulum is very large, and
is, as a food-store, very richly provided with albumin, fat, and other
nutritive matter (food-yolk); while the cytoplasm of the sperm-cell is
very small, and generally forms a vibrating lash, with which it moves
along and seeks the ovum.
In most of the plants the female and male cells are produced by the
same sprout, and in many of the lower animals by one and the same
person. This kind of hermaphrodism in "individuals of the second order"
is called monoclinism ("one-beddedness"). In many of the higher plants
(monœcic stocks) and most of the higher animals we have diclinism
("two-beddedness")--in other words, the one sprout or person has
only male, and the other sprout or person only female, organs--this
is gonochorism of individuals of the second order. Monoclinism is
generally associated with sedentary life (and often necessary for it),
and diclinism with free movement. Adaptation to parasitic habits also
favors monoclinism; thus, the crabs, for instance, are for the most
part gonochoristic individuals, but the creeping crabs (_cirripedia_),
which have adopted sedentary (and to an extent parasitic) habits, have
become hermaphrodites in consequence. Many intestinal parasites among
the lower animals (such as tape-worms, suctorial worms, wonder-snails),
which live isolated lives inside other animals, have to be
hermaphroditic and able to fertilise themselves if the species is to be
maintained. On the other hand, many hermaphroditic flowers, although
they have both sorts of sex-organs, are incapable of fertilizing
themselves and have to receive this from insect visitors which carry
the pollen from one flower to another.
Individuals of the third order, which we call stocks (_cormi_) in both
the plant and animal worlds, also exhibit varying features in the
sex-persons which compose them. When male and female diclinic sprouts
or persons are found side by side on the same stock, we call this
hermaphrodism of the cormi _monœcia_ ("one-housedness"); this is the
case with most of the siphonophora and some of the corals. _Diœcia_
("two-housedness") is less common: in this one stock has only male and
the other only female sprouts or persons, as in poplars and osiers,
most of the corals, and some of the siphonophora. The physiological
advantages of crossing--the union of sex-cells of different
individuals--favor progressive sex-division in the higher organisms.
A comparative study of the features of hermaphrodism and sex-division
in the plant and animal worlds teaches us that both forms of
sex-activity are often found in closely related organisms of one and
the same group, sometimes even in different individuals of the same
species. Thus, for instance, the oyster is usually gonochoristic, but
sometimes hermaphroditic; and so with many other mollusks, vermalia,
and articulata. Hence, the question often raised, which of the two
forms of sex-division is original, is hardly susceptible of a general
answer, or without relation to the stage of individuality and the place
in classification of the group under discussion. It is certain that in
many cases hermaphrodism represents the original feature; for instance,
in most of the lower plants and many of the stationary animals
(sponges, polyps, platodes, tunicates, etc.). Where we find exceptions
in these groups, they are of secondary origin. It is equally certain,
on the other hand, that in other cases the separation of the sexes is
the primitive arrangement; as in siphonophoræ, ctenophoræ, bryozoa,
cirripedia, and mollusks. In these cases the hermaphrodism is clearly
secondary in the sense that the hermaphrodites descend originally from
gonochorists.
It is only in a few sections of the lowest histona that the two kinds
of sex-cells arise without a definite location in different parts of
the simple tissue, as in a few groups of the lower algæ and in the
sponges. As a rule they are formed only at definite positions and in a
special layer of the tissue-body, and mostly in groups, in the shape
of sexual glands (_gonades_). These bear special names in different
groups of the histona. The female glands are called archegonia in
the cryptogams, _nucellus_ (formed from the macrosporangia of the
pteridophyta) in the phanerogams, and ovaries in the metazoa. The male
glands are called antheridia in the cryptogams, pollen-sacs (formed
from the microsporangia of the ferns) in the phanerogams, and testicles
(_as spermaria_) in the metazoa. In many cases, especially in aquatic
lower animals, the ovula (as products of the ovaries) are discharged
directly outward. But, in most of the higher organisms, special sexual
ducts (_gonoductus_) have been formed to conduct both kinds of the
gonocyta out of the organism.
While the two kinds of sexual glands are usually located in different
parts of the generating organism, there are, nevertheless, a few cases
in which the sex-cells are formed directly and together from one and
the same gland. These glands are called hermaphroditic glands. Such
structures are very notable in several highly differentiated groups
of the metazoa, and have clearly been developed from gonochoristic
structures in lower forms. The class of crested medusæ, or ribbed
medusæ (ctenophoræ), contains glasslike, sea-dwelling cnidaria of a
peculiar and complicated build, which probably descend from hydromedusæ
(or craspedota). But whereas the latter have very simple gonochoristic
structures (four or eight monosexual glands in the course of the
radial canals or in the gastric wall), in the ctenophoræ the eight
hermaphroditic canals run in a meridian arch from one pole of the
cucumber-shaped body to the other. Each canal corresponds to a ciliary
streamer, and forms ovaries at one border and testicles at the other;
and these are so arranged that the eight intercostal fields (the spaces
between the eight streamers) are alternately male and female. Still
more curious are the hermaphroditic glands of the highly organized,
land-dwelling, and air-breathing lung-snails (_pulmonata_), to which
our common garden snail (_arion_) and vineyard snail (_helix_) belong.
Here we have a hermaphroditic gland with a number of tubes, each of
which forms ovaries in its outer part and sperma in the inner. Still
the two kinds of sex-cells lead separately outward.
In most of the lower and aquatic histona both kinds of sex-cells,
when they are ripe, fall directly into the water, and come together
there. But in most of the higher, and especially the terrestrial,
organisms special exits or conducting canals have been formed for the
sex-products, the sexual ducts (_gonoductus_); in the metazoa the
female have the general name of oviducts and the male spermaducts (or
_vasa deferentia_). In the viviparous histona special canals serve
for the conveyance of the sperm to the ovum, which remains inside the
mother's body; such are the neck of the archegonium in the cryptogams,
the pistil in the phanerogams, and the vagina in the metazoa. At the
outer opening of these conducting canals special copulative organs are
developed, as a rule.
When the ejected sex-cells do not directly encounter each other (as in
many aquatic organisms), special structures have to be formed to convey
the fertilizing sperm from the male to the female body. This process of
copulation becomes important, as it is associated with characteristic
feelings of pleasure, which may cause extreme psychic excitement; as
sexual love it becomes, in man and the higher animals, one of the most
powerful springs of vital activity. In many of the higher animals
(namely, vertebrates, articulates, and mollusks) there are also formed
a number of glands and other auxiliary organs which co-operate in the
copulation.
The manifold and intimate relations which exist, in man and the higher
animals (especially vertebrates and articulates), between their sexual
life and their higher psychic activity, have given rise to plenty of
"wonders of life." Wilhelm Bölsche has so ably described them in his
famous and popular work, _The Life of Love in Nature_, that I need only
refer the reader to it. I will only mention the great significance of
what are called "secondary sexual characters." These characteristics
of one sex that are wanting in the other, and that are not directly
connected with the sexual organs--such as the man's beard, the woman's
breasts, the lion's mane, or the goat's horns--have also an æsthetic
interest; they have, as Darwin showed, been acquired by sexual
selection, as weapons of the male in the struggle for the female,
and vice versa. The feeling of beauty plays a great part in this,
especially in birds and insects; the beautiful colors and forms which
we admire in the male bird of paradise, the humming-bird, the pheasant,
the butterfly, etc., have been formed by sexual selection (_cf._ the
_History of Creation_).
In various groups of the histona the male sex has become superfluous
in the course of time; the ovula develop without the need of
fertilization. That is particularly the case in many of the platodes
(trematodes) and articulates (crustacea and insects). In the bees we
have the remarkable feature that it is only decided at the moment
of laying the egg whether it is to be fertilized or not; in the one
event a female and in the other a male bee is formed from it. When
Siebold proved at Munich these facts of miraculous conception in
various insects, he was visited by the Catholic archbishop of the
city, who expressed his gratification that there was now a scientific
explanation possible of the conception of the Virgin Mary. Siebold
had, unfortunately, to point out to him that the inference from the
parthenogenesis of the articulate to that of the vertebrate was not
valid, and that all mammals, like all other vertebrates, reproduce
exclusively from impregnated ova. We also find parthenogenesis
among the metaphyta, as in the _chara crinita_ among the algæ, the
_antennaria alpina_ and the _alchemilla vulgaris_ among the flowering
plants. We are, as yet, ignorant for the most part of the causes
of this lapse of fertilization. Some light has been thrown on it,
however, by recent chemical experiments (the effect of sugar and
other water-absorbing solutions), in which we have succeeded in
parthenogenetically developing unfertilized ova.
In the higher animals the complete maturity and development of
the specific form are requisite for reproduction, but in many of
the lower animals it has been observed recently that ovula and
sperm-cells are even formed by the younger specimens in the larva
stage. If impregnation takes place under these conditions, larvæ of
the same form are born. And when these larvæ have afterwards reached
maturity and reproduced in this form, we call the process _dissogony_
("double-generation"). It is found in many of the cnidaria, especially
the medusæ. But if larvæ propagate by unfertilized ova, and so
reproduce their kind parthenogenetically, the process is known as
_pædogenesis_ ("young-generation"). It is found particularly in the
platodes (trematodes) and some of the insects (larvæ of _cecidomyca_
and other flies).
In a large number of lower animals and plants sexual and asexual
generation regularly alternate. Among the protists we find this
alternation of generation in the sporozoa; among the metaphyta in the
mosses and ferns; and among the metazoa in the cnidaria, platodes,
tunicates, etc. Often the two generations differ considerably in shape
and degree of organization. Thus, in the mosses the asexual generation
is the spore-forming moss capsule (_sporogonium_), while the sexual is
the moss plant with stalk and leaves (_culmus_). In the case of the
ferns, on the other hand, the latter is spore-forming and monogenetic,
while the thallus-formed, simple, and small fore-germ (_prothallium_)
is sexually differentiated. In most of the cnidaria a small stationary
polyp is developed out of the ovum of the free-swimming medusa, and
this polyp, in turn, generates by budding medusæ, which reach sexual
maturity. In the tunicates (salpa) a sexual social form alternates with
an asexual solitary form; the chain-salpa of the former are smaller and
differently shaped than the large individual salpa of the latter, which
again generate chains by budding. This special form of metagenesis was
the first to be observed, as it was in 1819 by the poet Chamisso, when
he sailed round the world. In other cases (for instance, in the closely
related _doliolum_) a sexual generation alternates with two (or more)
neutral ones. The explanation of these various forms of alternating
generations is given in the laws of latent heredity (atavism), division
of labor, and metamorphosis, and especially by the biogenetic law.
While in real metagenesis (alternation of generations in the strict
sense) the asexual generation propagates by budding or spore-formation,
this is done parthenogenetically in the cognate process of
heterogenesis. This it is which, especially in many of the articulates,
causes an immense increase of the species in a short time. Among the
insects we have the leaf-lice (aphides), and among the crustacea the
water-fleas (daphnides), that propagate in great numbers during warm
weather by unfertilized "summer-ova." It is not until the autumn that
males appear and fertilize the large "winter-ova"; in the following
spring the first parthenogenetic generation issues from the winter
eggs. The two heterogenetic generations are very different in the
parasitic suctorial worms (trematodes). From the fertilized ovum of the
hermaphrodite distoma we get simply constructed nurses (pædogenetic
larvæ), inside which cercaria are generated from unfertilized ova;
these travel, and are afterwards converted (inside another animal) into
distoma once more.
I have given (_General Morphology_, chap, ii., p. 104) the name of
strophogenesis to the complicated process of cell-reproduction,
which we find in the ontogeny of most of the higher histona, both
phanerogams and cœlomaria. In these there is not a real alternation
of generations, as the multicellular tissue-forming organism develops
directly from the impregnated ovum. But the process resembles
metagenesis in so far as the ontogenetic construction consists itself
in a repeated division of the cells. Many generations of cells proceed
by cleavage from the one stem-cell (the impregnated ovum) before two
of these cells become sexually differentiated, and form a generation
of sexual cells. However, the essential difference consists in the
fact that all these generations of cells--in the body of both the
higher animals and the flowering plants--remain joined together as
parts of a single bion (a unified physiological individual); but in
the alternation of generations each group produced is made up of a
number of bionta, which live as independent forms--often so different
from each other that they were formerly thought to be animals of
separate classes, such as the polyps and medusæ. Hence we must not
describe the reproductive circle of the phanerogams as an alternation
of generations, although it has started from the fern (by abbreviated
heredity).
All simple forms of sexual reproduction without alternation of
generations are comprised under the title of _hypogenesis_. The
generative cycle proceeds from ovum to ovum in one and the same
bion or physiological individual. This form of development is usual
with most of the higher animals and plants; it may proceed with or
without metamorphosis. The younger forms which arise temporarily in
the latter case, and are distinguished from the sexually ripe form
by the possession of the provisional (and subsequently disappearing)
organs--larva organs (for instance, the tadpole or the pupa), are
comprised under the general head of larvæ.
As a rule, only organisms of the same species seem to have sexual
union and generate fertile progeny. This was formerly a rigid dogma,
and served the purpose of defining the loose idea of the species.
It was said: "When two animals or plants can have fertile offspring
they belong to the same real species." This principle, which once
afforded support to the dogma of the constancy of species, has long
been discarded. We now know by numbers of sound experiments that not
only two closely related species, but even two species of different
genera, may have sexual intercourse in certain circumstances, and that
the hybrids thus generated can have fertile offspring, either by union
among themselves or with one of the parents. However, the disposition
to hybridism varies considerably, and depends on the unknown laws of
sexual affinity. This sexual affinity must be based on the chemical
properties of the plasm of the copulating cells, but it seems to show
a good deal of vagueness in its effect. As a rule, hybrids exhibit a
combination of the features of both parents.
It has been proved by many recent experiments that hybrids have a more
powerful build and can reproduce more strongly than pure offspring,
whereas pure selection has generally in time an injurious effect. A
freshening by the introduction of new blood seems to be good from time
to time. Hence, it is just the reverse of what the former dogma of the
constancy of species affirmed. The question of hybridism has, generally
speaking, no value in defining the species. Probably many so-called
"true species," which have relatively constant features, are really
only permanent hybrids. This applies especially to lower sea-dwelling
animals, the sexual products of which are poured into the water and
swarm together in millions. As we know of various species of fishes,
crabs, sea-urchins, and vermalia, that their hybrids are very easily
produced and maintained by artificial impregnation, there is nothing to
prevent us from believing that such hybrids are also maintained in the
natural state.
The short survey we have made of the manifold varieties of reproduction
is sufficient to give an idea of the extraordinary wealth of this world
of wonders. When we go more closely into details we find hundreds of
other remarkable variations of the process on which the maintenance
of the species depends. But the most important point of all is the
fact that all the different forms of tocogony may be regarded as
connected links of a chain. The steps of this long ladder extend
uninterruptedly from the simple cell-division of the protists to the
monogony of the histona, and from this to the complicated amphigony
of the higher organisms. In the simplest case, the cell-cleavage of
the monera, propagation (by simple transverse division) is clearly
nothing more than transgressive growth. But even the preliminary
stage of sexual differentiation, the copulation of two equal cells
(_gameta_), is really nothing but a special form of growth. Then, when
the two gameta become unequal in the division of labor, when the larger
inert macrogameton stores up food in itself, and the smaller, mobile
microgameton swims in search of it, we have already expressed the
difference between the female ovum and the male sperm-cell. And in this
we have the most essential feature of sexual reproduction.
The reproduction of the organism is often regarded as a perfect mystery
of life, and as the vital function which most strikingly separates
the living from the lifeless. The error of this dualistic notion is
clear the moment one impartially considers the whole gradation of
forms of reproduction, from the simplest cell-division to the most
elaborate form of sexual generation, in phylogenetic connection. It is
obvious all through that transgressive growth is the starting-point
in the formation of new individuals. But the same must be said of
the multiplication of inorganic bodies--the cosmic bodies on the
larger scale, crystals on the smaller scale. When a rotating sun
passes a certain limit of growth by the constant accession of falling
meteorites, nebulous rings are detached at its equator by centrifugal
force, and form into new planets. Every inorganic crystal, too, has
a certain limit of individual growth (determined by its chemical and
molecular constitution). However much mother-water you add, this
is never passed, but new crystals (daughter-crystals) form on the
mother-crystal. In other words, growing crystals propagate.
XII
MOVEMENT
Mechanics as the science of motion (kinematics and
phoronomism)--Chemistry of vital movement--Active and passive
movements--Undulatory movement--Mechanism of imbibition--Autonomous
and reflex movements--Will and willing--Mixed movements--Movements
of growth--Direction of the vital movement--Direction of the
crystallizing force--Direction of cosmic motion--Movements of
protists--Amœboid, myophenous, hydrostatic, secretory, vibratory
movements: cilia and lashes--Movements of histona, metaphyta, and
metazoa--Locomotion of tissue animals: ciliary motion and muscular
movements--Muscles of the skin--Active and passive organs of
movement--Radiata, articulata, vertebrata, mammalia--Human movements.