In tracing the genealogy of any American
family it is often difficult or impossible to say
whether a certain branch is descended from John Oldworthy
or his cousin or second cousin. In the latter
cases to find the common ancestor we must go back to
the grandfather or great-grandfather. The same
difficulty, but greatly enhanced, meets us when we
try to make a genealogical tree of the animal kingdom.
Thus it seems altogether probable that all higher forms
are descended from an ancestor of the same general
structure and grade of organization as the turbellaria,
although probably free swimming, and hence with somewhat
different form and development, especially of the
muscular system. It seems to me altogether probable
that all, except possibly Mollusca, are descended
from a common ancestor closely resembling the schematic
worm last described. Some would, however, maintain
that they diverged rather earlier than even the turbellaria;
others after the schematic worm, if such ever existed.
As far as our argument is concerned it makes little
difference which of these views we adopt.
From our turbellaria, or possibly
from some even more primitive ancestor, many lines
diverged. And this was to be expected. The
coelenterata, as we saw in hydra, had developed rude
digestive and reproductive systems. The higher
groups of this kingdom had developed all, or nearly
all, the tissues used in building the bodies of higher
animals muscular, reproductive, connectile,
glandular, nervous, etc. But these are mostly
very diffuse. The muscular fibrils of a jelly-fish
are mostly isolated or parallel in bands, rarely in
compact well-defined bundles. The tissues have
generally not yet been moulded into compact masses
of definite form. There are as yet very few structures
to which we can give the name of organs. To form
organs and group them in a body of compact definite
form was the work pre-eminently of worms. The
material for the building was ready, but the architecture
of the bilateral animal was not even sketched.
And different worms were their own architects, untrammelled
by convention or heredity, hence they built very different,
sometimes almost fantastic, structures.
We must remember, too, the great age
of this group. They are present in highly modified
forms in the very oldest palaeozoic strata, and probably
therefore came into existence as the first traces of
continental areas were beginning to rise above the
primeval ocean. They are literally “older
than the hills.” They were exposed to a
host of rapidly changing conditions, very different
in different areas. This prepares us for the
fact that the worms represent a stage in animal life
corresponding fairly well to the Tower of Babel in
biblical history. The animal kingdom seems almost
to explode into a host of fragments. Our genealogical
tree fairly bristles with branches, but the branches
do not seem to form any regular whorls or spirals.
Few of them have developed into more than feeble growths.
They now contain generally but few species. Many
of them are largely or entirely parasitic, and in
connection with this mode of life have undergone modifications
and degeneration which make it exceedingly difficult
to decipher their descent or relationships.
Four of these branches have reached
great prominence in numbers and importance. One
or two others were formerly equally numerous and have
since become almost extinct; so the brachiopoda, which
have been almost entirely replaced by mollusks.
The same may very possibly be true of others.
For of the amount of extinction of larger groups we
have generally but an exceedingly faint conception.
Indeed in this respect the worms have been well compared
to the relics which fill the shelves of one of our
grandmother’s china-closets.
The four great branches are the echinoderms,
mollusks, articulates, and vertebrates. The echinoderms,
including starfishes, sea-urchins, and others straggled
early from the great army. We know as yet almost
nothing of their history; when deciphered it will be
as strange as any romance. The vertebrates are
of course the most important line, as including the
ancestors of man. But we must take a little glance
at mollusks, including our clams, snails, and cuttle-fishes;
and at the articulates, including annelids and culminating
in insects. The molluscan and articulate lines,
though divergent, are of great importance to us as
throwing a certain amount of light on vertebrate development;
and still more as showing how a certain line of development
may seem, and at first really be, advantageous, and
still lead to degeneration, or at best to but partial
success.
When we compare the forms which represent
fairly well the direction of development of these
three lines, a snail or a clam with an insect and
a fish, we find clearly, I think, that the fundamental
anatomical difference lies in the skeleton; and that
this resulted from, and almost irrevocably fixed,
certain habits of life.
We may picture to ourselves the primitive
ancestor of mollusks as a worm having the short and
broad form of the turbellaria, but much thicker or
deeper vertically. A fuller description can be
found in the “Encyclopædia Britannica,”
Art., Mollusca. It was hemi-ovoid in form.
It had apparently the perivisceral cavity and nephridia
of the schematic worm, and a circulatory system.
In this latter respect it stood higher than any form
which we have yet studied. Its nervous system
also was rather more advanced. It had apparently
already taken to a creeping mode of life and the muscles
of its ventral surface were strongly developed, while
its exposed and far less muscular dorsal surface was
protected by a cap-like shell covering the most important
internal organs. But the integument of the whole
dorsal surface was, as is not uncommon in invertebrates,
hardening by the deposition of carbonate of lime in
the integument. And this in time increased to
such an extent as to replace the primitive, probably
horny, shell.
Into the anatomy of this animal or
of its descendants we have no time to enter, for here
we must be very brief. We have already noticed
that the most important viscera were lodged safely
under the shell. And as these increased in size
or were crowded upward by the muscles of the creeping
disk, their portion of the body grew upward in the
form of a “visceral hump.” Apparently
the animal could not increase much in length and retain
the advantage of the protection of the shell; and
the shell was the dominating structure. It had
entered upon a defensive campaign. Motion, slow
at the outset, became more difficult, and the protection
of the shell therefore all the more necessary.
The shell increased in size and weight and motion
became almost impossible. The snail represents
the average result of the experiment. It can
crawl, but that is about all; it is neither swift
nor energetic. Even the earthworm can outcrawl
it. It has feelers and eyes, and is thus better
provided with sense-organs than almost any worm.
It has a supra-oesophageal ganglion of fair size.
The clams and oysters show even more
clearly what we might call the logical results of
molluscan structure. They increased the shell
until it formed two heavy “valves” hanging
down on each side of the body and completely enclosing
it. They became almost sessile, living generally
buried in the mud and gaining their food, consisting
mostly of minute particles of organic matter, by means
of currents created by cilia covering the large curtain-like
gills. Their muscular system disappeared except
in the ploughshare-shaped “foot” used
mostly for burrowing, and in the muscles for closing
the shell. That portion of the body which corresponds
to the head of the snail practically aborted with
nearly all the sense-organs. The nervous system
degenerated and became reduced to a rudiment.
They had given up locomotion, had withdrawn, so to
speak, from the world; all the sense they needed was
just enough to distinguish the particles of food as
they swept past the mouth in the current of water.
They have an abundance of food, and “wax fat.”
The clam is so completely protected by his shell and
the mud that he has little to fear from enemies.
They have increased and multiplied and filled the mud.
“Requiescat in pace.”
But zooelogy has its tragedies as
well as human history. Let us turn to the development
of a third molluscan line terminating in the cuttle-fishes.
The ancestors of these cephalopods, although still
possessed of a shell and a high visceral hump, regained
the swimming life. First, apparently, by means
of fins, and then by a simple but very effective use
of a current of water, they acquired an often rapid
locomotion. The highest forms gave up the purely
defensive campaign, developed a powerful beak, led
a life like that of the old Norse pirates, and were
for a time the rulers and terrors of the sea.
With their more rapid locomotion the supra-oesophageal
ganglion reached a higher degree of development, and
it was served by sense-organs of great efficiency.
They reduced the external shell, and succeeded, in
the highest forms, of almost ridding themselves of
this burden and encumbrance. Traces of it remain
in the squids, but transformed into an internal quill-like,
supporting, not defensive, skeleton. They have
retraced the downward steps of their ancestors as
far as they could. And the high development of
their supra-oesophageal ganglion and sense-organs,
and their powerful jaws and arms, or tentacles, show
to what good purpose they have struggled. But
the struggle was in vain, as far as the supremacy
of the animal kingdom was concerned. Their ancestors
had taken a course which rendered it impossible for
their descendants to reach the goal. Their progress
became ever slower. They were entirely and hopelessly
beaten by the vertebrates. They struggled hard,
but too late.
The history of mollusks is full of
interest. They show clearly how intimately nervous
development is connected with the use of the locomotive
organs. The snail crept, and slightly increased
its nervous system and sense-organs. The clam
almost lost them in connection with its stationary
life. The cephalopods were exceedingly active,
developed, therefore, keen sense-organs and a very
large and complicated supra-oesophagal ganglion, which
we might almost call a brain.
The articulate series consists of
two groups of animals. The higher group includes
the crabs, spiders, thousand-legs, and finally the
insects, and forms the kingdom of arthropoda.
The lower members are still usually reckoned as worms,
and are included under the annelids. Of these
our common earthworm is a good example, and near them
belong the leeches. But the marine annelids, of
which nereis, or a clam-worm, is a good example, are
more typical. They are often quite large, a foot
or even more in length. They are composed of
many, often several hundred, rings or segments.
Between these the body-wall is thin, so that the segments
move easily upon each other, and thus the animal can
creep or writhe.
These segments are very much alike
except the first two and the last. If we examine
one from the middle of the body we shall find its
structure very much like that of our schematic worm.
Outside we find a very thin, horny cuticle, secreted
by the layer of cells just beneath it, the hypodermis.
Beneath the skin we find a thin layer of transverse
muscles, and then four heavy bands of longitudinal
muscles. These latter have been grouped in the
four quadrants, a much more effective arrangement
than the cylindrical layer of the schematic worm.
Furthermore, the animal has on each segment a pair
of fin-like projections, stiffened with bristles, the
parapodia. These are moved by special muscles
and form effective organs of creeping.
Within the muscles is the perivisceral
cavity, and in its central axis the intestine, segmented
like the body-wall. The reproductive organs are
formed from patches of the lining of the perivisceral
cavity, and the reproductive elements, when fully developed,
fall into the perivisceral fluid and are carried out
by nephridia, just such as we found in the schematic
worm. Beside the perivisceral cavity and its
fluid there is a special circulatory system. This
consists mainly of one long tube above the intestine
and a second below, with often several smaller parallel
tubes. Transverse vessels run from these to all
parts of the body. The dorsal tube pulsates and
thus acts as a heart. The surface of the body
no longer suffices to gather oxygen, hence we find
special feathery gills on the parapodia. But
these gills are merely expanded portions of the body
wall, arranged so as to offer the greatest possible
amount of surface where the capillaries of the blood
system can be almost immediately in contact with the
surrounding water.
The nervous system consists of a large
supra-oesophageal ganglion in the first segment; then
of a chain of ganglia, one to each segment, on the
ventral side of the body. With one ganglion in
each segment there is far more controlling, perceptive,
ganglionic material than in lower worms. Furthermore
the supra-oesophageal ganglion is relieved of a large
part of the direct control of the muscles of each
segment, and is becoming more a centre of control
and perception for the body as a whole. It is
more like our brain, commander-in-chief, the other
ganglia constituting its staff. The sense-organs
have improved greatly. There are tentacles and
otolith vesicles as very delicate organs of feeling,
or possibly of hearing also.
But the annelids were probably the
first animals to develop an eye capable of forming
an image of external objects. The importance of
this organ in the pursuit of food or the escape from
enemies can scarcely be over-estimated. The lining
of the mouth and pharynx can be protruded as a proboscis,
and drawn back by powerful muscles, and is armed with
two or more horny claws. Eyes and claws gave them
a great advantage over their not quite blind but really
visionless and comparatively defenceless neighbors,
and they must have wrought terrible extinction of
lower and older forms. But while we cannot over-estimate
the importance of these eyes, we can easily exaggerate
their perfectness. They were of short range, fitted
for seeing objects only a few inches distant, and
the image was very imperfect in detail. But the
plan or fundamental scheme of these eyes is correct
and capable of indefinitely greater development than
the organs of touch or smell, perhaps greater even
than the otolith vesicle.
And the reflex influence of the eye
on the brain was the greatest advantage of all.
Hitherto with feeble muscles and sense-organs it has
hardly paid the animal to devote more material to building
a larger brain. It was better to build more muscle.
But now with stronger muscles at its command, and
better sense-organs to report to it, every grain of
added brain material is beginning to be worth ten
devoted to muscle. The muscular system will still
continue to develop, but the brain has begun an almost
endless march of progress. The eye becomes of
continually increasing advantage and importance because
it has a capable brain to use it; and brain is a more
and more profitable investment, because it is served
by an ever-improving eye.
The annelid had hit upon a most advantageous
line of development, which led ultimately to the insect.
The study of the insect will show us clearly the advantages
and defects of the annelid plan. First of all,
the insect, like the mollusk, has an external skeleton.
But the skeleton of the mollusk was purely protective,
a hindrance to locomotion. That of the insect
is still somewhat protective, but is mainly, almost
purely, locomotive. It is never allowed to become
so heavy as to interfere with locomotion. In the
second place, the insect has three body regions, having
each its own special functions or work. And one
of these is a head. The annelid had two anterior
segments differing from those of the rest of the body;
these may, perhaps, be considered as the foreshadowings
of a structure not yet realized; they can only by
courtesy be called a head. Thirdly, the insect
has legs. The annelid had fin-like parapodia,
approaching the legs of insects about as closely as
the fins of a fish approach the legs of a mammal.
The reproductive and digestive systems, while somewhat
improved, are not very markedly higher than those
of annelids. The excretory system has more work
to perform and reaches a rather higher development.
But in these organs there is no great
or striking change; the time for marked and rapid
development of the digestive and reproductive systems
has gone by. Material can be more profitably invested
in brain or muscle. Air is carried to all parts
of the body by a special system of air-sacks and tubes.
This is a very advantageous structure for small animals
with an external skeleton. In very large animals,
or where the skeleton is internal, it would hardly
be practicable; the risk of compression of the tubes
at some point, and of thus cutting off the air-supply
of some portion of the body, would be altogether too
great.
The circulatory system is very poor.
It consists practically only of a heart, which drives
the blood in an irregular circulation between the
other organs of the body much as with a syringe you
might keep up a system of currents in a bowl of water.
But the rapidity of the flow of the blood in our bodies
is mainly to furnish a supply of oxygen to the organs.
A tea-spoonful of blood can carry a fair amount of
dissolved solid nutriment like sugar, it can carry
at each round but a very little gas like oxygen.
Hence the blood must make its rounds rapidly, carrying
but a little oxygen at each circuit. But in the
insect the blood conveys only the dissolved solid
nutriment, the food; hence a comparatively irregular
circulation answers all purposes.
The skeleton is a thickening of the
horny cuticle of the annelid on the surface of each
segment. The horny cylinder surrounding each
segment is composed of several pieces, and on the abdomen
these are united by flexible, infolded membranes.
This allows the increase in the size of the segment
corresponding to the varying size of the digestive
and reproductive systems. In this part of the
body the skeletal ring of each segment is joined to
that of the segments before and behind it in the same
manner. But in other parts of the body we shall
find the skeletal pieces of each segment and the rings
of successive segments fused in one plate of mail.
The legs are the parapodia of annelids carried to
a vastly higher development. They are slender
and jointed, and yet often very powerful. A large
portion of the muscular system of the body is attached
to these appendages.
But the insect has also jaws.
The annelid had teeth or claws attached to the proboscis.
But true jaws are something quite different.
They always develop by modifying some other organ.
In the insect they are modified legs. This is
shown first by their embryonic development. But
the king- or horseshoe-crab has still no true jaws,
but uses the upper joints of its legs for chewing.
There are primitively three pairs of jaws of various
forms for the different kinds of food of different
species or higher groups. But some of them may
disappear and the others be greatly modified into
awls for piercing, or a tube for sucking honey.
Into the wonderful transformations of these modified
legs we cannot enter.
The muscles are no longer arranged
to form a sack as in annelids. Transverse muscles,
running parallel to the unyielding plates of chitin
or horn could accomplish nothing. They have largely
disappeared. The work of locomotion has been transferred
from the trunk to the legs.
The abdomen of the insect is as clearly
composed of distinct segments as the body of the annelid.
Of these there are perhaps typically eleven.
The thorax is composed of three segments, distinct
in the lowest forms, fused in the highest. This
fusion of segments in the thorax of the highest forms
furnishes a very firm framework for the attachment
of wings and muscles. These wings are a new development,
and how they arose is still a question. But they
give the insect the capability of exceedingly rapid
locomotion.
The three pairs of jaws, modified
legs, in the rear half of the head show that this
portion is composed of three segments. For only
one pair of legs is ever developed on a single segment.
Embryology has shown that the portion of the head
in front of the mouth is also composed of three segments.
Possibly between the prae- and post-oral portions
still another segment should be included, making a
total of seven in the head. The head has thus
been formed by drawing forward segments from the trunk,
and fusing them successively with the first or primitive
head segment. This is difficult to conceive of
in the fully developed insect, where the boundary
between head and thorax is very sharp. But the
ancestors of insects looked more like thousand-legs
or centipedes, and here head and thorax are much less
distinct. But in the annelid the mouth is on the
second segment; here it is on the fourth. It
has evidently travelled backward. That the mouth
of an animal can migrate seems at first impossible,
but if we had time to examine the embryology of annelids
and insects, it would no longer appear inconceivable
or improbable. And its backward migration brought
it among the legs which were grasping and chewing
the food. And in vertebrates the mouth has changed
its position, though not in exactly the same way.
Our present mouth is probably not at all the mouth
of the primitive ancestor of vertebrates. Thus
in the insect three segments have fused around the
mouth, and three, possibly four, in front of it.
This makes a head worthy of the name. The ganglia
of the three post-oral segments, which bear the jaws,
have fused in one compound ganglion innervating the
mouth and jaws. Those of the three prae-oral
segments have fused to form a brain. Eyes are
well developed, giving images sometimes accurate in
detail, sometimes very rude. Ears are not uncommon.
The sense of smell is often keen.
Perhaps the greatest advance of the
insect is its adaptation to land life. This gives
it a larger supply of oxygen than any aquatic animal
could ever obtain. This itself stimulates every
function, and all the work of the body goes on more
energetically. Then the heat produced is conducted
off far less rapidly than in aquatic forms. Water
is a good conductor of heat, and nearly all aquatic
animals are cold-blooded. The few which are warm-blooded
are protected by a thick layer of non-conducting fat.
In all land animals, even when cold-blooded, the work
of the different systems is aided by the longer retention
of the heat in the body.
Let us recapitulate. The schematic
worm had a body composed of two concentric tubes.
The outer was composed of the muscles of the body
covered by the protective integument. The inner
tube was the alimentary canal with its special muscles.
Between these two was the perivisceral cavity, filled
with nutritive fluid, lymph, and furnishing a safe
lodging-place for the more delicate viscera. It
represented fairly the trunk of higher animals.
The annelid added segmentation, and
thus greater freedom of motion by the parapodia.
But the segments were still practically alike.
In the insect division of labor took place, that is,
each group of segments was allotted its own special
work; and these groups of segments were modified in
structure to best suit the performance of this part
of the work of the body. The abdomen was least
modified and its eleven segments were devoted to digestion,
reproduction, and excretion the old vegetative
functions. Three segments were united in the
thorax; all their energy was turned to locomotion,
and the insect became thus an exceedingly active,
swift animal. The third body-region, the head,
includes six segments, of which three surrounded the
mouth and furnished the jaws, while two more were
crowded or drawn forward in order that their ganglia
might be added to the old supraoesophageal ganglion
and form a brain. It is interesting to note that
a form, peripatus, still exists which stands almost
midway between annelids and insects and has only four
segments in the head. The formation of the head
was thus a gradual process, one segment being added
after another.
In the turbellaria the dominant functions
were digestion and reproduction, and their organs
composed almost the whole body. Here only eleven
segments at most are devoted to these functions, and
nine in head and thorax to locomotion and brain.
Head and thorax have increased steadily in importance,
while the abdomen has decreased as steadily in number
of segments. And the brain is increasing thus
rapidly because there are now muscles and sense-organs
of sufficient power to make such a brain of value.
And this brain perceives not only objects and qualities,
but invisible relations between these, and this is
an advance amounting to a revolution. It remembers,
and uses its recollections. It is capable of
learning a little by experience and observation.
The A, B, C of thinking was probably learned long
before the insect’s time, and the bee shows
a fair amount of intelligence.
The line of development which the
insect followed was comparatively easy and its course
probably rapid. Certain crustacea, aquatic arthropoda,
are among the oldest fossils, and it is possible that
insects lived on the land before the first fish swam
in the sea. They had fine structure and powers;
and yet during the later geologic periods they have
scarcely advanced a step, and are now apparently at
a standstill. They ran splendidly for a time,
and then fell out of the race. What hindered
and stopped them?
One vital defect in their whole plan
of organization is evident. The external skeleton
is admirably suited to animals of small size, but
only to these. In larger animals living on land
it would have to be made so heavy as to be unwieldy
and no longer economical. Their mode of breathing
also is fitted only for animals of small size having
an external skeleton. Whatever may be our explanation
the fact remains that insects are always small.
This is in itself a disadvantage. Very small
animals cannot keep up a constant high temperature
unless the surrounding air is warm, for their radiating
surface is too large in comparison with their heat-producing
mass. At the first approach of even cool weather
they become chilled and sluggish, and must hibernate
or die. They are conformed to but a limited range
of environment in temperature.
But small size is, as a rule, accompanied
by an even greater disadvantage. It seems to
be almost always correlated with short life.
Why this is so, or how, we do not know. There
are exceptions; a crow lives as long as a man; or
would, if allowed to. But, as a rule, the length
of an animal’s days is roughly proportional to
the size of its body. And the insect is, as a
rule, very short-lived. It lives for a few days
or weeks, or even months, but rarely outlasts the
year. It has time to learn but little by experience.
The same experience must be passed, the same emergency
arise and be met, over and over again during the lifetime
of the same individual if the animal is to learn thereby.
And intelligence is based upon experience. Hence
insects can and do possess but a low grade of intelligence.
But instinct is in many cases habit fixed by heredity
and improved by selection. The rapid recurrence
of successive generations was exceedingly favorable
to the development of instincts, but very unfavorable
to intelligence. Insects are instinctive, the
highest vertebrates intelligent. The future can
never belong to a tiny animal governed by instincts.
Mollusks and insects have both failed to reach the
goal; another plan of structure than theirs must be
sought if the animal kingdom is to have a future.
The future belonged to the vertebrate.
To begin with less characteristic organs the digestive
system is much like that of the annelid or schematic
worm, but with greatly increased glandular and absorptive
surfaces. The present mouth of nearly all vertebrates
is probably not primitive. It is almost certainly
one of the gill-slits of some old ancestor of fish,
such as now are used to discharge the water which
is used for respiration. The jaws are modified
branchial arches or the cartilaginous or bony rods
which in our present fish support the fringe of gills.
These have formed a pair of exceedingly effective
and powerful jaws. The reproductive system holds
still to the old type and shows little if any improvement.
The excretory organs, kidneys, are composed primitively
of nephridial tubes like those of the schematic worm
or annelid, but immensely increased in number, modified,
and improved in certain very important particulars.
The muscles in simplest forms are composed of heavy
longitudinal bands, especially developed toward the
dorsal surface of the body to the right and left of
the axial skeleton. Locomotion was produced by
lashing the tail right and left, as still in fish.
There is improvement in all these organs, except perhaps
the reproductive, but nothing very new or striking.
The great improvement from this time on was not to
be sought in the vegetative organs, or even directly
to any great extent in muscles.
The new and characteristic organ was
not the vertebral column, or series of vertebrae,
or backbone, from which the kingdom has derived its
name. This was a later production. The primitive
skeleton was the notochord, still appearing in the
embryos of all vertebrates and persisting throughout
life in fish. This is an elastic rod of cartilage,
lying just beneath the spinal marrow or nerve-cord,
which runs backward from the brain. The nerve-centres
are therefore here all dorsal, and the notochord or
skeleton lies between these and the digestive or alimentary
canal. The skeleton of the clam or snail is purely
protective and a hindrance to locomotion. That
of the insect is almost purely locomotive, but external,
that of the vertebrate purely locomotive and internal.
It does not lie outside even of the nervous system,
although this system especially required, and was
worthy of, protection. It does not protect even
the brain; the skull of vertebrates is an after-thought.
It is almost the deepest seated of all organs.
But lying in the central axis of the body it furnishes
the very best possible attachment for muscles.
Around this primitive notochord was a layer of connectile
tissue which later gave rise to the vertebrae forming
our backbone.
The nervous system on the dorsal surface
of the notochord consists of the brain in the head
and the spinal marrow running down the back.
The brain of all except the very lowest vertebrates
consists of four portions: 1. The cerebrum,
or cerebral lobes, or simply “forebrain,”
the seat of consciousness, thought, and will, and from
which no nerves proceed. Whether the primitive
vertebrate had any cerebrum is still uncertai.
The mid-brain, which sends nerves to the eyes, and
in this respect reminds us of the brain of insects.
Its anterior portion appears from embryology to be
very primitiv. The small brain, or cerebellum,
which in all higher forms is the centre for co-ordination
of the motions of the bod. The medulla, which
controls especially the internal organs. The spinal
marrow, or that portion of the nervous system which
lies outside of the head, is at the same time a great
nerve-trunk and a centre for reflex action of the
muscles of the body. But the development of these
distinct portions and the division of labor between
them must have been a long and gradual process.
We have every reason to believe that
here, as in insects, the head has been formed by annexation
of segments from the rump and the fusion of their
nervous matter with that of the brain. But here,
instead of only three segments, from nine to fourteen
have been fused in the head to furnish the material
for the brain. Notochord and backbone may be
the most striking and apparent characteristic of vertebrates,
but their predominant characteristic is brain.
On this system they lavished material, giving it from
three to four times as much as any lower or earlier
group had done. They very early set apart the
cerebral lobes to be the commander-in-chief and centre
of control for all other nerve-centres. To this
all report, and from it all directly or indirectly
receive orders. It can say to every other organ
in the body, “Starve that I may live.”
It is the seat of thought and will. The other
portions of the brain report to it what they have
gathered of vision or sound; it explains the vision
or song or parable. It is relieved as far as
possible from all lower and routine work that it may
think and remember and govern. The vertebrate
built for mind, not neglecting the body.
Every trait of vertebrates is a promise
of a great future. Its internal skeleton gives
it the possibility of large size. This gave it
in time the victory in the struggle with its competitors,
as to whether it should eat or be eaten. It is
vigorous and powerful, for all its organs are at the
best. It gives the possibility of later, on land,
becoming warm-blooded, i.e., of maintaining
a constant high temperature. It is thus resistant
to climate and hardship. In time its descendants
will face the arctic winter as well as the heat of
the tropics.
But it has started on the road which
leads to mind. The greater size is correlated
with longer life. The lessons of experience come
to it over and over again, and it can and must learn
them. It is the intelligent, remembering, thinking
type. The insect had begun to peer into the world
of invisible and intangible relations, the vertebrate
will some day see them. This much is prophecied
in his very structure. He must be heir to an
indefinite future.
You have probably noticed that the
vertebrate differs greatly from all his predecessors.
The gulf between him and them is indeed wide and deep.
His origin and ancestry are yet far from certain.
But an attempt to decipher his past history, though
it may lead to no sure conclusions, will yet be of
use to us. Practically all aquatic vertebrates
lead a swimming life, neither sessile nor creeping.
The embryonic development of our appendages leads
to the same conclusion. We must never forget
that the embryonic development of the individual recapitulates
briefly the history of the development of the race.
Now the legs and arms, or fore- and hind-legs, of
higher vertebrates and the corresponding paired fins
of fish develop in the embryo as portions of a long
ridge extending from front to rear of the side of
the body.
This justifies the inference that
the primitive vertebrate ancestor had a pair of long
fins running along the sides of the body, but bending
slightly downward toward the rear so as to meet one
another and continue as a single caudal fin behind
the anal opening. Such fins, like the feathers
of an arrow, could be useful only to keep the animal
“on an even keel” as it was forced through
the water by the lateral sweeps of the tail.
They would have been useless for creeping.
But there is another piece of evidence
that he was a free swimming form. All vertebrates
breathe by gills or lungs, and these are modified
portions of the digestive system, of the walls of the
oesophagus, from which even the lung is an embryonic
outgrowth. Now practically all invertebrates
breathe through modified portions of the integument
or outer surface of the body, and their gills are
merely expansions of this. In the annelid they
are projections of the parapodia, in the mollusk expansions
of the skin, where the foot or creeping sole joins
the body. Why did the vertebrate take a new and
strange, and, at first sight, disadvantageous mode
of breathing? There must have been some good
reason for this. The most natural explanation
would seem to be that he had no projections on his
outer surface which could develop into gills, and farther,
that he could not afford to have any. Now projections
on the lower portion of the sides of the body would
be an advantage in creeping, but a hindrance in any
such mode of swimming as we have described, or indeed
in any mode of writhing through the water.
Furthermore, if he lived, not a creeping
life on the bottom, but swimming in the water above,
he would have to live almost entirely on microscopic
animals and embryos; and these would be most easily
captured by a current of water brought in at the mouth.
The whole branchial apparatus in its simplest forms
would seem to be an apparatus for sifting out the
microscopic particles of food and only later a purely
respiratory apparatus. Moreover, we have seen
that the parapodia of annelids naturally point to
the development of an external skeleton, for their
muscles are already a part of the external body-wall
and attached to the already existing horny cuticle.
The logical goal of their development was the insect.
Now I do not wish to conceal from
you that many good zooelogists believe that the vertebrate
is descended from annelids; but for this and other
reasons such a descent appears to me very improbable.
It would seem far more natural to derive the vertebrate
from some free swimming form like the schematic worm,
whose largest nerve-cord lay on the dorsal surface
because its branches ran to heavy muscles much used
in swimming. Later the other nerve-cords degenerated,
for such a degeneration of nerve-cords is not at all
impossible or improbable. “No thoroughfare”
is often written across paths previously followed
by blood or nervous impulses, when other paths have
been found more economical or effective.
But where did the notochord come from?
I do not know. It always forms in the embryo
out of the entoderm or layer which becomes the lining
of the intestine. Now this is a very peculiar
origin for cartilage, and the notochord is a very
strange cartilage even if we have not made a mistake
in calling it cartilage at all. My best guess
would be that it is simply a thickened portion of the
upper median surface of the intestine to keep the
“balls” of digesting nutriment or other
hard particles in the intestine from “grinding”
against the nerve-cord as they are crowded along in
the process of digestion. Once started its elasticity
would be a great aid in swimming.
Professor Brooks has called attention
to the fact that the higher a group stands in development,
the longer its ancestors have maintained a swimming
life. Thus we have noticed that the sponges were
the first to settle; then a little later the mass of
the coelenterates followed their example. But
the etenophora, the nearest relatives of bilateral
animals, have remained free swimming. Then the
flat worms and mollusks took to a creeping mode of
life, while the annelids and vertebrates still swam.
Then the annelids settled to the bottom and crept,
and all their descendants remained creeping forms.
The vertebrates alone remained swimming, and probably
neither they nor their descendants ever crept until
they emerged on the land, or as amphibia were preparing
for land life. If this be true, it is a fact
worthy of our most careful consideration. The
swimming life would appear to be neither as easy nor
as economical as the creeping. It is certainly
hard to believe that food would not have been obtained
with less effort and in greater abundance at the bottom
than in the water above. The swimming life gave
rise to higher and stronger forms; but did its maintenance
give immediate advantage in the struggle for existence?
This is an exceedingly interesting and important question,
and demands most careful consideration. But we
shall be better prepared to answer it in a future
lecture.
The period of development of mollusks,
articulates, and vertebrates, is really one.
They developed to a certain extent contemporaneously.
The development of vertebrates was slow, and they were
the last to appear on the stage of geological history.
You must all have noticed that development,
during this period, takes on a much more hopeful form
than during that described in the last chapter.
Then digestion and reproduction were dominant.
Now muscle is of the greatest importance. If
this fails of development, as in mollusks, the group
is doomed to degeneration or at best stagnation.
But we have seen the dawn of a still higher function.
In insects and vertebrates the brain is becoming of
importance, and absorbing more and more material.
This is the promise of something vastly higher and
better. Better sense-organs are appearing, fitted
to aid in a wider perception of more distant objects.
The vertebrate has discovered the right path; though
a long journey still lies before it. The night
is far spent, the day is at hand.