Form and Function
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E. S. (Edward Stuart) Russell >> Form and Function
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"Nature seems to operate always according to an original and general
plan, from which she departs with regret and whose traces we come
across everywhere" (Vicq d'Azyr, quoted by Flourens, _Mem. Acad.
Sei._, XXIII., p. xxxvi.).
Peter Camper (1722-1789), we are told by Goethe himself in his
_Osteologie_, was convinced of the unity of plan holding throughout
Vertebrates; he compared in particular the brain of fishes with the
brain of man.
The idea of the unity of plan had not yet become limited and defined
as a strictly scientific theory; it was an idea common to philosophy,
to ordinary thought, and to anatomical science. We find it expressed
by Herder (who perhaps got it from Kant) in his _Ideen sur Philosophie
der Geschichte der Menschheit_ (1784), and it is possible that Goethe
became impressed with the importance of the idea through his
conversations with Herder. Be that as it may, it is certain that
Goethe sought for the intermaxillaries in man only because he was
firmly convinced that the skeleton in all the higher animals was built
upon one common plan and that accordingly bones such as the
intermaxillaries, found well developed in some animals, must also be
found in man. The idea was not drawn from the facts, but the facts
were interpreted and even sought for in the light of the idea. "I
eagerly worked upon a general osteological scheme, and had accordingly
to assume that all the separate parts of the structure, in detail as
in the whole, must be discoverable in all animals, because on this
supposition is built the already long begun science of comparative
anatomy."[72]
The principle comes to clear expression in his _Erster Entwurf einer
allgemeinen Einleitung in die vergleichende Anatomie_ (1795).[73] He
writes:--"On this account an attempt is here made to arrive at an
anatomical type, a general picture in which the forms of all animals
are contained in potentia, and by means of which we can describe each
animal in an invariable order."[74] His aim is to discover a general
scheme of the constant in organic parts, a scheme into which all
animals will fit equally well, and no animal better than the rest.
When we remember that the type to which anatomists before him had,
consciously or unconsciously, referred all other structure was man
himself, we see that in seeking after an abstract generalised type
Goethe was reaching out to a new conception. The fact that only the
structure of man and the higher animals was at all well-known in his
time led Goethe to think that his general Typus would hold for the
lower animals as well, though it was to be arrived at primarily from a
study of the higher animals. All he could assert of the entire animal
kingdom was that all animals agreed in having a head, a middle part,
and an end part, with their characteristic organs, and that
accordingly they might, in this respect at least, be reduced to one
common Typus. Goethe's knowledge of the lower animals was not
extensive.
Though Goethe did not work out a criterion of the homology of parts
with any great clearness, he had an inkling of the principle later
developed by E. Geoffroy St Hilaire, and called by him the "Principle
of Connections." According to this principle, the homology of a part
is determined by its position relative to other parts. Goethe
expresses it thus:--"On the other hand the most constant factor is the
position in which the bone is invariably found, and the function to
which it is adapted in the organic edifice."[75] But from this sentence
it is not clear that Goethe understood the principle as one of form
independent of function, for he seems to consider that the homology of
an organ is partly determined by the function which it performs for
the whole. He wavers between the purely formal or morphological
interpretation of the principle of connections and the functional. We
find him in the additions to the _Entwurf_ (1796), saying:--"We must
take into consideration not merely the spatial relations of the parts,
but also their living reciprocal influence, their dependence upon and
action on one another." [76] But in seeking for the intermaxillary bone
in man he was guided by its position relative to the maxillaries--it
must be the bone between the anterior ends of the maxillaries, a bone
whose limits are indicated in the adult only by surface grooves.
As a matter of fact Goethe's morphological views are neither very
clearly expressed nor very consistent. This comes out in his treatment
of the relation between structure and function. Sometimes he takes the
view that structure determines function. "The parts of the animal," he
writes, "their reciprocal forms, their relations, their particular
properties determine the life and habits of the creature."[77] We are
not to explain, he says, the tusks of the _Babirussa_ by their
possible use, but we must ask how it comes to have tusks. In the same
way we must not suppose that a bull has horns in order to gore, but we
must investigate the process by which it comes to have horns to gore
with. This is the rigorous morphological view. On the other hand he
admits elsewhere that function may influence form. Apparently he did
not work out his ideas on this point to logical clearness, and Radl[78]
is probably correct in saying that the following quotation with its
double assertion represents most nearly Goethe's position:--
"Also bestimmt die Gestalt die Lebensweise des Thieres, Und die Weise
zu leben, sie wirkt auf alle Gestalten Maechtig zurueck."[79]
His best piece of purely morphological work was his theory of the
metamorphosis of plants. Stripped of its vaguer elements, and of the
crude attempt to explain differences in the character of plant organs
by differences in the degree of "refinement" of the sap supplied to
them, the theory is that stem-leaves, sepals, petals, and stamens are
all identical members or appendages. These appendages differ from one
another only in shape and in degree of expansion, stem-leaves being
expanded, sepals contracted, petals expanded, and so on alternately.
It is equally correct to call a stamen a contracted petal, and a petal
an expanded stamen, for no one of the organs is the type of the
others, but all equally are varieties of a single abstract
plant-appendage.
What Goethe considered he had proved for the appendages of plants he
extended to all living things. Every living thing is a complex of
living independent beings, which "der Idee, der Anlage nach," are the
same, but in appearance may be the same or similar, different or
unlike.[80] Not only is there a primordial animal and a primordial
plant, schematic forms to which all separate species are referable,
but the parts of each are themselves units, which "der Idee nach," are
identical _inter se_. This fantasy can hardly be taken seriously as a
scientific theory; it seems, however, to have been what guided Goethe
in his "discovery" of the vertebral nature of the skull. Just as the
fore limb can be homologised with the hind limb, so, reasoning by
analogy, the skull should be capable of being homologised with the
vertebrae. To what ludicrous extremes this doctrine of the repetition
of parts within the organism was pushed we shall see when we consider
the theories of the German transcendentalists of the early nineteenth
century.
Though Goethe's morphological views were lacking in definiteness he
hit upon one or two ideas which proved useful. Thus he enunciated the
"law of balance" long before Etienne Geoffroy St Hilaire, the law
"that to no part can anything be added, without something being taken
away from another part, and _vice versa_."[81] He saw, too, what a help
to the interpretation of adult structure the study of the embryo would
be, for many bones which are fused in the adult are separate in the
embryo.[82] This also was a point to which the later transcendentalists
gave considerable attention.
So far we have spoken of Goethe as if he were merely the prophet of
formal morphology; we have pointed out how he brought to clear
expression the morphological principle implicit in the idea of unity
of type, and how he seized upon some important guiding ideas, such as
the principle of connections. But Goethe was not a formalist, and he
was very far from the static conception of life which is at the base
of pure morphology. His interest was not in _Gestalt_ or fixed form,
_Bildung_ or form change. He saw that _Gestalt_ was but a momentary
phase of _Bildung_, and could be considered apart and in itself only
by an abstraction fatal to all understanding of the living thing.
Mephistopheles scoffs at the scholars who would explain a living
creature by anatomising it:
"Dann hat er die Theile in seiner Hand,
Fehlt leider! nur das geistige Band."[83]
Goethe kept clear of this mistake; he knew that the artist comes
nearer to the truth than the analyst.
In the fragment entitled _Bildung und Umbildung organischer Naturen_
(1807), introductory to a reprint of his paper on the "Metamorphosis
of Plants," we get an exposition of his general views on living
things. He points out there how we try to understand things by
separating them into their parts. We can, it is true, resolve the
organism into its structural elements, but we cannot recompose it or
endow it with life by joining up the parts. Hence we require some
other means of understanding it. "In all ages even among scientific
men there can be discerned a yearning to apprehend the living form as
such, to grasp the connection of their external visible parts, to
interpret them as indications of the inner activity, and so, in a
certain measure, to master the whole conceptually." This science which
should discover the inner meaning of organic _Bildung_ is called
Morphology.[84] In Morphology we should not speak of _Gestalt_ or fixed
form, or if we do we should understand by it only a momentary phase of
_Bildung_. Form is of interest not in itself but only as the
manifestation of the inner activity of the living being. Over
development, he says elsewhere, there presides a formative force, a
_bildende Kraft_ or _Bildungstrieb_, which works out the idea of the
organism. Living things, in his view of them, strive to manifest an
idea. They are Nature's works of art--and so, incidentally, they
require an artist to interpret them.
This profound conception of the nature of life is applied not only to
the growing changing individual but also to the whole changing world
of organisms. They are all manifestations of a living shaping power
which moulds them. This shaping power, immanent in all life, is
conceived to work according to a general plan, and so we get an
explanation of the fact that living things seem simply varieties of
one common type.
"If we once recognise," says Goethe, "that the creative spirit brings
into being and shapes the evolution of the more perfect organic
creatures according to a general scheme, is it altogether impossible
to represent this original plan if not to the senses at least to the
mind...?"[85]
Such an interpretation of the unity of plan reaches perhaps beyond the
bounds of science.
[70] _See_ Kohlbrugge, "Hist. krit. Studien ueber Goethe
als Naturforscher," _Zool. Annalen._ v., 1913, pp.
83-231.
[71] Or re-discovered, according to Kohlbrugge.
[72] Cotta ed., vol. ix., p. 448.
[73] "First Draft of a General Introduction to
Comparative Anatomy."
[74] Cotta ed., ix., p. 463.
[75] Cotta ed., p. 478.
[76] _Loc. cit._, p. 491.
[77] _Entwurf_, Cotta ed., ix., p. 465.
[78] _Geschichte der biologischen Theorien_, i., p. 266.
[79] "So the form determines the manner of life of the
animal, and the manner of life in its turn reacts
powerfully upon all forms."
[80] _Bildung und Umbildung organischer Naturen_, 1807.
[81] Cotta ed., ix., p. 466.
[82] _Loc. cit._, pp. 474-5.
[83] Then he has all the parts within his hand, excepting
only, sad to say, the living bond.
[84] Goethe was the inventor of the word.
[85] Cotta ed., ix., p. 490.
CHAPTER V
ETIENNE GEOFFROY SAINT-HILAIRE
E. Geoffrey made an experiment, unsuccessful but instructive. He tried
to found a science of pure morphology; he failed: his failure showed,
once and for all, that a pure morphology of organic forms is
impracticable.
Already, in 1796, in one of his earliest memoirs,[86] Geoffroy was
guided by the idea that Nature has formed all living things upon one
plan. Organs which seem anomalous are merely modifications of the
normal; the trunk of an elephant is formed by the excessively
prolonged nostrils, the horn of a rhinoceros is simply a mass of
adhering hairs. In general, however varied their form, all organs are
simply variations of a common scheme; Nature employs no new organs.
Organs which are rudimentary, such as the clavicles in the ostrich and
the nictitating membrane in man, bear witness to the unity of plan. In
this Geoffroy goes no further than his predecessors. They too had
recognised homologies of organs; they too had interpreted rudimentary
organs as vestiges of an original plan.
In a series of papers published in 1807, Geoffroy took a further step,
and sought to establish homologies which were not obvious--homologies,
too, not so much of organs as of parts.
These memoirs (published in the _Annales du Museum d'Histoire
naturelle_, vols. ix. and x., 1807) dealt with the homology between
the bones of the pectoral fin and girdle in fish and the bones of the
arm and shoulder-girdle in higher Vertebrates, with the homologies of
the bones of the sternum, and with the determination of the pieces of
the skull, particularly in the crocodile. All Geoffroy's morphological
doctrine is found in them, but for the full expression of his views we
must take his chief work, the _Philosophie anatomique_, particularly
the first volume (1818). This volume contains, beside the important
"Discours preliminaire" and "Introduction" which we shall presently
consider in detail, five memoirs, which deal with the various bones
connected with the respiratory organs in fishes (the bones of the
operculum, of the hyoid, of the branchial arches, of the pectoral
girdle), and seek to discover their homologies with corresponding
bones in air-breathing Vertebrates.
"Can the organisation of vertebrated animals be referred to one
uniform type?" This is the question with which the _Philosophie
anatomique_ opens, the question to which the whole book is an answer.
But is it not generally acknowledged by naturalists that Vertebrates
are built upon one uniform plan, that, for instance, the fore limb may
be modified for running, climbing, swimming, or flying, yet the
arrangement of the bones remain the same? How else could there be a
"natural method" of classification?[87]
But the homologies so drawn repose upon a vague and confused feeling for
likenesses; they are not based upon an explicit principle. What general
principle can be applied? "Now it is evident that the sole general
principle one can apply is given by the position, the relations, and the
dependencies of the parts, that is to say, by what I name and include
under the term of _connections_." For instance, the part known as the
hand in man and generally as the fore foot in other Vertebrates, is the
fourth part in order in the anterior member, and its homologue can
always be recognised by this fact of its connections (p. xxvi.). The
principle of connections serves as a guide in tracing an organ through
all its functional transformations, for "an organ can be deteriorated,
atrophied, annihilated, but not transposed" (p. xxx.).
It is this principle which enables one to follow out in detail the
further fundamental conception that in every Vertebrate there are found
the same "organic materials," or units of construction. This conception,
which Geoffroy calls the _Theorie des analogues_ (p. xxxii.), is clearly
one part of the old idea of the unity of type; it teaches the _unity of
composition_ of organic beings, while the _Principe des connexions_ adds
the _unity of plan_.
Both conceptions are logically implicit in the vague notion of unity of
type; Geoffroy disengaged them, and pushed each to its logical extreme.
Most of the ordinary homologies of structure in air-breathing
Vertebrates have already been seized, he continues, for they are more or
less obvious, and many intermediate states exist (p. xxxiv.). But
ordinary methods of comparison fail when the attempt is made to
homologise the structure of fishes with that of air-breathing
Vertebrates, for the homologies are anything but obvious and no
intermediate organs are found.
Most air-breathing Vertebrates have a larynx, a trachea, and bronchi,
which are absent in fish; and fish have many parts which seem to be
absent in higher Vertebrates. But apply the "Theory of Analogues"; it
teaches that there can be no organ peculiar to fish and not found in
other Vertebrates; apply the "Principle of Connections," it will show
which organs are homologous in the two types (p. xxxv.).
Comparative anatomists, with few exceptions, had hitherto taken man as
the type, and referred all structure to his; Geoffroy's principles led
him to give preference to no one animal in particular, but to seize upon
each part in the species in which it reaches the maximum of its
development (p. xxxvi.). He is thus led to refer all structures to a
generalised abstract type. In this abstract type each organ exists at
the maximum of its development, each organ shows all its potentialities
realised. In a way, therefore, this type, this abstraction, gives the
scheme of the possible transformations of each organ.
It is true Geoffroy does not refer to this "Archetype" in so many words,
but it must always have been vaguely present in his mind. He has this
idea in his head when he says in one of his later works, "There is,
philosophically speaking, only a single animal."[88] The "single animal"
is simply the generalised type.
Having laid down his two principles Geoffroy goes on to apply them to
the difficult case of the comparison of the skeleton of fish with the
skeleton of the higher Vertebrates. "My present task is to demonstrate
that there is no part of the bony framework of fishes that cannot find
its analogue in the other vertebrated animals."[89] It seems at first
sight that many bones are peculiar to fish, formed expressly for
performing the functions which fish do not share with higher animals.
These are the bones connected with respiration--the operculum, the
branchiostegal rays, the branchial arches, and others. That the peculiar
bones should be connected with the respiratory functions is only
natural, for the contrast between fish and higher Vertebrates is
essentially a contrast between water-breathing and air-breathing
animals. Considering first the general form of the skeleton in fish, we
are met at once with a difficulty; there is no obvious homologue in
fishes of the neck, the trunk, and the abdomen of higher animals. What
apparently corresponds to the trunk is in fishes crowded close up under
the head. But, after all, it is not of the essence of the vertebrate
type to have the trunk and the abdomen attached at definite and
invariable distances along the vertebral column--that is a notion
surviving from the anatomy which made man its type. The "trunk" differs
in position according to the class, in quadrupeds, birds, and fishes (p.
9). Now, says Geoffroy, allow me this one hypothesis, that the trunk
with its organs can, as it were, move bodily along the vertebral column,
so as to be found in one class near the front end of the vertebral
column, in another about the middle, and in a third near the end, then I
can show you in detail that the constituent parts of this trunk are
found in all classes to be invariably in the same positions relatively
to one another (p. 10). It is important to note this hypothesis of a
"metastasis" which Geoffroy makes, for it is the key to the
understanding of many of the far-fetched homologies which he tries to
establish. It is, of course, clear that this hypothesis is in formal
contradiction with his principal hypothesis of the invariability of
connections, and that he, so to speak, gets a hold on his fish to apply
his principle of connections only by admitting at the very outset an
exception to his primary principle. A further application of the
hypothesis of metastasis will be noticed below in connection with the
determination of the sternum of fishes. We note here an interpretation
of the first metastasis in terms of functional adaptation. "The constant
and violent action of the tail, if it does not go so far as actually to
displace and move forward the internal organs, at least fits in well
with an arrangement in which the organs are so disposed" (p. 99).
The first memoir deals with the homologies of the opercular bones.
Geoffroy considers that the external opening of the ear corresponds to
the external opening of the gill-chamber, which lies between the
operculum and the pectoral girdle. The ear communicates with the buccal
cavity by the Eustachian tube, so does the branchial chamber by means of
the gill-slits. The auditory chamber of higher Vertebrates is,
therefore, the homologue of the branchial chamber in fish; the opercular
bones in fish and the ossicles of the ear in other Vertebrates stand in
close relation to this chamber; therefore the opercular bones are the
homologues of the ossicles of the ear, the interoperculum corresponding
to the malleus, the suboperculum to the lenticular, the minute lower
part of the suboperculum to the incus, the operculum to the stapes, and
the pre-operculum to the tympanic ring. In making these particular
determinations Geoffroy professes to be led by his principle of
connections. The pre-operculum has, he says, the same connections with
neighbouring bones as the tympanic bone in other Vertebrates, and the
other pieces of the gill-cover are homologised with particular
ear-ossicles according to the order in which they stand to one another.
The second memoir in the book deals with the sternum, and affords a very
good example of Geoffroy's method of dealing with the facts of
structure. We shall omit here any detailed reference to the other three
memoirs, which deal with the hyoid, with the branchial arches and the
structures which correspond in air-breathing Vertebrates, and with the
bones of the shoulder-girdle.
In the memoir on the sternum Geoffroy's first care is to arrive at a
definition of what a sternum is. He defines it partly by its functions,
partly by its connections, as the system of bones which covers and
protects the thorax, and gives attachment to certain groups of muscles.
The most highly developed sternum (according to this definition) is the
plastron of the tortoise, whose structure it dominates (p. 103). It is
important, therefore, to determine of how many bones the plastron is
composed, since the full number of elementary parts of which an organ is
composed is best seen when the organ is at the maximum of its
development. There are nine bones in the plastron of the tortoise. "The
conclusion to be drawn from this is that every sternum, provided that it
is not inhibited in its development by some obstacle, is composed of
_nine elementary parts_" (p. 105). These nine bones are in Geoffroy's
nomenclature, the episternals, the hyosternals, the hyposternals, the
xiphisternals, which are all paired bones, and the entosternal, which is
unpaired. The arrangement of them is in the tortoise:--
Episternal---------------------------Episternal
|\__ __/|
| \__ __/ |
| \__ __/ |
| \__ Entosternal __/ |
| __/ \__ |
| __/ \__ |
| __/ \__ |
|/ \|
Hyosternal Hyosternal
| |
| |
| |
| |
Hyposternal-------------------------Hyposternal
| |
| |
| |
| |
Xiphisternal------------------------Xiphisternal.
The articulations in the tortoise are indicated by the connecting
lines. Geoffroy tries to show that the sternum in other animals is
composed of these nine bones, or at least of a certain number of them,
always in the same invariable relative positions. Thus in birds the
sternum consists of five pieces, of a huge keeled entosternal, and of
two "annexes" on either side, which are the hyo-and hyposternals.
These are separate only in young birds. Occasionally, especially in
young birds, rudiments of episternals and xiphisternals also occur.
The minuteness of the episternals and the xiphisternals may be
attributed to the gigantic size of the entosternal, in accordance with
the _Loi de balancement_. In the other air-breathing Vertebrates the
nine sternal elements can according to Geoffroy be discovered without
great difficulty. But when we come to the determination of the sternum
in fishes, difficulties abound, which Geoffroy solves in the following
way. He points out that between the clavicles (_cleithra_) and the
hyoid bone (_basihyal_) in fishes there is a long median bone
(_urohyal_) which is attached in front by two strong tendons to the
horns of the hyoid and is free behind (see Fig. 1). Gouan (1720) had
seen in this bone the homologue of the sternum. Geoffroy adopts this
view, but considers that this bone alone cannot represent the whole
sternum. He finds the representatives of other bones of the sternum in
the large bones (_epihyal_ and _ceratohyal_, or the two pieces of the
_ceratohyal_) which are comprised in the hyoid arch. But he is
immediately met by the difficulty that this complex of bones is
situated in front of the pectoral girdle, whereas the sternum in
higher Vertebrates lies behind the pectoral girdle. He reflects,
however, that the gills of fish, situated in front of the clavicles,
are merely the lungs under another name. The gills have become shifted
forward by a metastasis similar to that which brought the whole
thoracic organs far forward in fish. This being so, their supporting
elements, the sternum and the ribs, must have moved with them, and are
hence to be found in front of the pectoral girdle.
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