SigPhi · Charles Darwin

The Origin of Species by Means of Natural Selection

Page 16 of 36

prominent, recurved teeth, like those of the Merganser (a member of the same family), serving for the widely different purpose of securing live fish.

Returning to the whales. The Hyperoodon bidens is destitute of true teeth in an efficient condition, but its palate is roughened, according to Lacepede, with small unequal, hard points of horn. There is, therefore, nothing improbable in supposing that some early Cetacean form was provided with similar points of horn on the palate, but rather more regularly placed, and which, like the knobs on the beak of the goose, aided it in seizing or tearing its food. If so, it will hardly be denied that the points might have been converted through variation and natural selection into lamellæ as well-developed as those of the Egyptian goose, in which case they would have been used both for seizing objects and for sifting the water; then into lamellæ like those of the domestic duck; and so onward, until they became as well constructed as those of the shoveller, in which case they would have served exclusively as a sifting apparatus. From this stage, in which the lamellæ would be two-thirds of the length of the plates of baleen in the Balænoptera rostrata, gradations, which may be observed in still-existing Cetaceans, lead us onward to the enormous plates of baleen in the Greenland whale. Nor is there the least reason to doubt that each step in this scale might have been as serviceable to certain ancient Cetaceans, with the functions of the parts slowly changing during the progress of development, as are the gradations in the beaks of the different existing members of the duck-family. We should bear in mind that each species of duck is subjected to a severe struggle for existence, and that the structure of every part of its frame must be well adapted to its conditions of life.

The Pleuronectidæ, or Flat-fish, are remarkable for their asymmetrical bodies. They rest on one side—in the greater number of species on the left, but in some on the right side; and occasionally reversed adult specimens occur. The lower, or resting-surface, resembles at first sight the ventral surface of an ordinary fish; it is of a white colour, less developed in many ways than the upper side, with the lateral fins often of smaller size. But the eyes offer the most remarkable peculiarity; for they are both placed on the upper side of the head.

During early youth, however, they stand opposite to each other, and the whole body is then symmetrical, with both sides equally coloured. Soon the eye proper to the lower side begins to glide slowly round the head to the upper side; but does not pass right through the skull, as was formerly thought to be the case. It is obvious that unless the lower eye did thus travel round, it could not be used by the fish while lying in its habitual position on one side. The lower eye would, also, have been liable to be abraded by the sandy bottom. That the Pleuronectidæ are admirably adapted by their flattened and asymmetrical structure for their habits of life, is manifest from several species, such as soles, flounders, &c., being extremely common. The chief advantages thus gained seem to be protection from their enemies, and facility for feeding on the ground. The different members, however, of the family present, as Schiödte remarks, “a long series of forms exhibiting a gradual transition from Hippoglossus pinguis, which does not in any considerable degree alter the shape in which it leaves the ovum, to the soles, which are entirely thrown to one side.”

Mr. Mivart has taken up this case, and remarks that a sudden spontaneous transformation in the position of the eyes is hardly conceivable, in which I quite agree with him. He then adds: “If the transit was gradual, then how such transit of one eye a minute fraction of the journey towards the other side of the head could benefit the individual is, indeed, far from clear. It seems, even, that such an incipient transformation must rather have been injurious.” But he might have found an answer to this objection in the excellent observations published in 1867 by Malm. The Pleuronectidæ, while very young and still symmetrical, with their eyes standing on opposite sides of the head, cannot long retain a vertical position, owing to the excessive depth of their bodies, the small size of their lateral fins, and to their being destitute of a swimbladder. Hence, soon growing tired, they fall to the bottom on one side. While thus at rest they often twist, as Malm observed, the lower eye upward, to see above them; and they do this so vigorously that the eye is pressed hard against the upper part of the orbit. The forehead between the eyes consequently becomes, as could be plainly seen, temporarily contracted in breadth.

On one occasion Malm saw a young fish raise and depress the lower eye through an angular distance of about seventy degrees.

We should remember that the skull at this early age is cartilaginous and flexible, so that it readily yields to muscular action. It is also known with the higher animals, even after early youth, that the skull yields and is altered in shape, if the skin or muscles be permanently contracted through disease or some accident. With long-eared rabbits, if one ear flops forward and downward, its weight drags forward all the bones of the skull on the same side, of which I have given a figure.

Malm states that the newly-hatched young of perches, salmon, and several other symmetrical fishes, have the habit of occasionally resting on one side at the bottom; and he has observed that they often then strain their lower eyes so as to look upward; and their skulls are thus rendered rather crooked. These fishes, however, are soon able to hold themselves in a vertical position, and no permanent effect is thus produced. With the Pleuronectidæ, on the other hand, the older they grow the more habitually they rest on one side, owing to the increasing flatness of their bodies, and a permanent effect is thus produced on the form of the head, and on the position of the eyes. Judging from analogy, the tendency to distortion would no doubt be increased through the principle of inheritance. Schiödte believes, in opposition to some other naturalists, that the Pleuronectidæ are not quite symmetrical even in the embryo; and if this be so, we could understand how it is that certain species, while young, habitually fall over and rest on the left side, and other species on the right side. Malm adds, in confirmation of the above view, that the adult Trachypterus arcticus, which is not a member of the Pleuronectidæ, rests on its left side at the bottom, and swims diagonally through the water; and in this fish, the two sides of the head are said to be somewhat dissimilar. Our great authority on Fishes, Dr. Günther, concludes his abstract of Malm’s paper, by remarking that “the author gives a very simple explanation of the abnormal condition of the Pleuronectoids.”

We thus see that the first stages of the transit of the eye from one side of the head to the other, which Mr. Mivart considers would be injurious, may be attributed to the habit, no doubt beneficial to the individual and to the species, of endeavouring to look upward with both eyes, while resting on one side at the bottom. We may also attribute to the inherited effects of use the fact of the mouth in several kinds of flat-fish being bent towards the lower surface, with the jaw bones stronger and more effective on this, the eyeless side of the head, than on the other, for the sake, as Dr. Traquair supposes, of feeding with ease on the ground. Disuse, on the other hand, will account for the less developed condition of the whole inferior half of the body, including the lateral fins; though Yarrel thinks that the reduced size of these fins is advantageous to the fish, as “there is so much less room for their action than with the larger fins above.” Perhaps the lesser number of teeth in the proportion of four to seven in the upper halves of the two jaws of the plaice, to twenty-five to thirty in the lower halves, may likewise be accounted for by disuse. From the colourless state of the ventral surface of most fishes and of many other animals, we may reasonably suppose that the absence of colour in flat-fish on the side, whether it be the right or left, which is under-most, is due to the exclusion of light. But it cannot be supposed that the peculiar speckled appearance of the upper side of the sole, so like the sandy bed of the sea, or the power in some species, as recently shown by Pouchet, of changing their colour in accordance with the surrounding surface, or the presence of bony tubercles on the upper side of the turbot, are due to the action of the light. Here natural selection has probably come into play, as well as in adapting the general shape of the body of these fishes, and many other peculiarities, to their habits of life. We should keep in mind, as I have before insisted, that the inherited effects of the increased use of parts, and perhaps of their disuse, will be strengthened by natural selection. For all spontaneous variations in the right direction will thus be preserved; as will those individuals which inherit in the highest degree the effects of the increased and beneficial use of any part. How much to attribute in each particular case to the effects of use, and how much to natural selection, it seems impossible to decide.

I may give another instance of a structure which apparently owes its origin exclusively to use or habit. The extremity of the tail in some American monkeys has been converted into a wonderfully perfect prehensile organ, and serves as a fifth hand. A reviewer, who agrees with Mr. Mivart in every detail, remarks on this structure: “It is impossible to believe that in any number of ages the first slight incipient tendency to grasp could preserve the lives of the individuals possessing it, or favour their chance of having and of rearing offspring.” But there is no necessity for any such belief. Habit, and this almost implies that some benefit great or small is thus derived, would in all probability suffice for the work. Brehm saw the young of an African monkey (Cercopithecus) clinging to the under surface of their mother by their hands, and at the same time they hooked their little tails round that of their mother. Professor Henslow kept in confinement some harvest mice (Mus messorius) which do not possess a structurally prehensive tail; but he frequently observed that they curled their tails round the branches of a bush placed in the cage, and thus aided themselves in climbing. I have received an analogous account from Dr. Günther, who has seen a mouse thus suspend itself. If the harvest mouse had been more strictly arboreal, it would perhaps have had its tail rendered structurally prehensile, as is the case with some members of the same order. Why Cercopithecus, considering its habits while young, has not become thus provided, it would be difficult to say. It is, however, possible that the long tail of this monkey may be of more service to it as a balancing organ in making its prodigious leaps, than as a prehensile organ.

The mammary glands are common to the whole class of mammals, and are indispensable for their existence; they must, therefore, have been developed at an extremely remote period, and we can know nothing positively about their manner of development. Mr. Mivart asks: “Is it conceivable that the young of any animal was ever saved from destruction by accidentally sucking a drop of scarcely nutritious fluid from an accidentally hypertrophied cutaneous gland of its mother? And even if one was so, what chance was there of the perpetuation of such a variation?” But the case is not here put fairly. It is admitted by most evolutionists that mammals are descended from a marsupial form; and if so, the mammary glands will have been at first developed within the marsupial sack. In the case of the fish (Hippocampus) the eggs are hatched, and the young are reared for a time, within a sack of this nature; and an American naturalist, Mr. Lockwood, believes from what he has seen of the development of the young, that they are nourished by a secretion from the cutaneous glands of the sack. Now, with the early progenitors of mammals, almost before they deserved to be thus designated, is it not at least possible that the young might have been similarly nourished? And in this case, the individuals which secreted a fluid, in some degree or manner the most nutritious, so as to partake of the nature of milk, would in the long run have reared a larger number of well-nourished offspring, than would the individuals which secreted a poorer fluid; and thus the cutaneous glands, which are the homologues of the mammary glands, would have been improved or rendered more effective. It accords with the widely extended principle of specialisation, that the glands over a certain space of the sack should have become more highly developed than the remainder; and they would then have formed a breast, but at first without a nipple, as we see in the Ornithorhyncus, at the base of the mammalian series. Through what agency the glands over a certain space became more highly specialised than the others, I will not pretend to decide, whether in part through compensation of growth, the effects of use, or of natural selection.

The development of the mammary glands would have been of no service, and could not have been affected through natural selection, unless the young at the same time were able to partake of the secretion. There is no greater difficulty in understanding how young mammals have instinctively learned to suck the breast, than in understanding how unhatched chickens have learned to break the egg-shell by tapping against it with their specially adapted beaks; or how a few hours after leaving the shell they have learned to pick up grains of food. In such cases the most probable solution seems to be, that the habit was at first acquired by practice at a more advanced age, and afterwards transmitted to the offspring at an earlier age. But the young kangaroo is said not to suck, only to cling to the nipple of its mother, who has the power of injecting milk into the mouth of her helpless, half-formed offspring. On this head Mr. Mivart remarks: “Did no special provision exist, the young one must infallibly be choked by the intrusion of the milk into the wind-pipe. But there _is_ a special provision. The larynx is so elongated that it rises up into the posterior end of the nasal passage, and is thus enabled to give free entrance to the air for the lungs, while the milk passes harmlessly on each side of this elongated larynx, and so safely attains the gullet behind it.” Mr. Mivart then asks how did natural selection remove in the adult kangaroo (and in most other mammals, on the assumption that they are descended from a marsupial form), “this at least perfectly innocent and harmless structure?” It may be suggested in answer that the voice, which is certainly of high importance to many animals, could hardly have been used with full force as long as the larynx entered the nasal passage; and Professor Flower has suggested to me that this structure would have greatly interfered with an animal swallowing solid food.

We will now turn for a short space to the lower divisions of the animal kingdom. The Echinodermata (star-fishes, sea-urchins, &c.) are furnished with remarkable organs, called pedicellariæ, which consist, when well developed, of a tridactyle forceps—that is, of one formed of three serrated arms, neatly fitting together and placed on the summit of a flexible stem, moved by muscles. These forceps can seize firmly hold of any object; and Alexander Agassiz has seen an Echinus or sea-urchin rapidly passing particles of excrement from forceps to forceps down certain lines of its body, in order that its shell should not be fouled. But there is no doubt that besides removing dirt of all kinds, they subserve other functions; and one of these apparently is defence.

With respect to these organs, Mr. Mivart, as on so many previous occasions, asks: “What would be the utility of the _first rudimentary beginnings_ of such structures, and how could such insipient buddings have ever preserved the life of a single Echinus?” He adds, “not even the _sudden_ development of the snapping action would have been beneficial without the freely movable stalk, nor could the latter have been efficient without the snapping jaws, yet no minute, nearly indefinite variations could simultaneously evolve these complex co-ordinations of structure; to deny this seems to do no less than to affirm a startling paradox.” Paradoxical as this may appear to Mr.

Mivart, tridactyle forcepses, immovably fixed at the base, but capable of a snapping action, certainly exist on some star-fishes; and this is intelligible if they serve, at least in part, as a means of defence.

Mr. Agassiz, to whose great kindness I am indebted for much information on the subject, informs me that there are other star-fishes, in which one of the three arms of the forceps is reduced to a support for the other two; and again, other genera in which the third arm is completely lost. In Echinoneus, the shell is described by M. Perrier as bearing two kinds of pedicellariæ, one resembling those of Echinus, and the other those of Spatangus; and such cases are always interesting as affording the means of apparently sudden transitions, through the abortion of one of the two states of an organ.

With respect to the steps by which these curious organs have been evolved, Mr. Agassiz infers from his own researches and those of Mr.

Müller, that both in star-fishes and sea-urchins the pedicellariæ must undoubtedly be looked at as modified spines. This may be inferred from their manner of development in the individual, as well as from a long and perfect series of gradations in different species and genera, from simple granules to ordinary spines, to perfect tridactyle pedicellariæ.

The gradation extends even to the manner in which ordinary spines and the pedicellariæ, with their supporting calcareous rods, are articulated to the shell. In certain genera of star-fishes, “the very combinations needed to show that the pedicellariæ are only modified branching spines” may be found. Thus we have fixed spines, with three equi-distant, serrated, movable branches, articulated to near their bases; and higher up, on the same spine, three other movable branches.

Now when the latter arise from the summit of a spine they form, in fact, a rude tridactyle pedicellariæ, and such may be seen on the same spine together with the three lower branches. In this case the identity in nature between the arms of the pedicellariæ and the movable branches of a spine, is unmistakable. It is generally admitted that the ordinary spines serve as a protection; and if so, there can be no reason to doubt that those furnished with serrated and movable branches likewise serve for the same purpose; and they would thus serve still more effectively as soon as by meeting together they acted as a prehensile or snapping apparatus. Thus every gradation, from an ordinary fixed spine to a fixed pedicellariæ, would be of service.

In certain genera of star-fishes these organs, instead of being fixed or borne on an immovable support, are placed on the summit of a flexible and muscular, though short, stem; and in this case they probably subserve some additional function besides defence. In the sea-urchins the steps can be followed by which a fixed spine becomes articulated to the shell, and is thus rendered movable. I wish I had space here to give a fuller abstract of Mr. Agassiz’s interesting observations on the development of the pedicellariæ. All possible gradations, as he adds, may likewise be found between the pedicellariæ of the star-fishes and the hooks of the Ophiurians, another group of the Echinodermata; and again between the pedicellariæ of sea-urchins and the anchors of the Holothuriæ, also belonging to the same great class.

Certain compound animals, or zoophytes, as they have been termed, namely the Polyzoa, are provided with curious organs called avicularia.

These differ much in structure in the different species. In their most perfect condition they curiously resemble the head and beak of a vulture in miniature, seated on a neck and capable of movement, as is likewise the lower jaw or mandible. In one species observed by me, all the avicularia on the same branch often moved simultaneously backwards and forwards, with the lower jaw widely open, through an angle of about 90 degrees, in the course of five seconds; and their movement caused the whole polyzoary to tremble. When the jaws are touched with a needle they seize it so firmly that the branch can thus be shaken.

Mr. Mivart adduces this case, chiefly on account of the supposed difficulty of organs, namely the avicularia of the Polyzoa and the pedicellariæ of the Echinodermata, which he considers as “essentially similar,” having been developed through natural selection in widely distinct divisions of the animal kingdom. But, as far as structure is concerned, I can see no similarity between tridactyle pedicellariæ and avicularia. The latter resembles somewhat more closely the chelæ or pincers of Crustaceans; and Mr. Mivart might have adduced with equal appropriateness this resemblance as a special difficulty, or even their resemblance to the head and beak of a bird. The avicularia are believed by Mr. Busk, Dr. Smitt and Dr. Nitsche—naturalists who have carefully studied this group—to be homologous with the zooids and their cells which compose the zoophyte, the movable lip or lid of the cell corresponding with the lower and movable mandible of the avicularium.

Mr. Busk, however, does not know of any gradations now existing between a zooid and an avicularium. It is therefore impossible to conjecture by what serviceable gradations the one could have been converted into the other, but it by no means follows from this that such gradations have not existed.

As the chelæ of Crustaceans resemble in some degree the avicularia of Polyzoa, both serving as pincers, it may be worth while to show that with the former a long series of serviceable gradations still exists.

In the first and simplest stage, the terminal segment of a limb shuts down either on the square summit of the broad penultimate segment, or against one whole side, and is thus enabled to catch hold of an object, but the limb still serves as an organ of locomotion. We next find one corner of the broad penultimate segment slightly prominent, sometimes furnished with irregular teeth, and against these the terminal segment shuts down. By an increase in the size of this projection, with its shape, as well as that of the terminal segment, slightly modified and improved, the pincers are rendered more and more perfect, until we have at last an instrument as efficient as the chelæ of a lobster. And all these gradations can be actually traced.

Besides the avicularia, the polyzoa possess curious organs called vibracula. These generally consist of long bristles, capable of movement and easily excited. In one species examined by me the vibracula were slightly curved and serrated along the outer margin, and all of them on the same polyzoary often moved simultaneously; so that, acting like long oars, they swept a branch rapidly across the object-glass of my microscope. When a branch was placed on its face, the vibracula became entangled, and they made violent efforts to free themselves. They are supposed to serve as a defence, and may be seen, as Mr. Busk remarks, “to sweep slowly and carefully over the surface of the polyzoary, removing what might be noxious to the delicate inhabitants of the cells when their tentacula are protruded.” The avicularia, like the vibracula, probably serve for defence, but they also catch and kill small living animals, which, it is believed, are afterwards swept by the currents within reach of the tentacula of the zooids. Some species are provided with avicularia and vibracula, some with avicularia alone and a few with vibracula alone.

It is not easy to imagine two objects more widely different in appearance than a bristle or vibraculum, and an avicularium like the head of a bird; yet they are almost certainly homologous and have been developed from the same common source, namely a zooid with its cell.

Hence, we can understand how it is that these organs graduate in some cases, as I am informed by Mr. Busk, into each other. Thus, with the avicularia of several species of Lepralia, the movable mandible is so much produced and is so like a bristle that the presence of the upper or fixed beak alone serves to determine its avicularian nature. The vibracula may have been directly developed from the lips of the cells, without having passed through the avicularian stage; but it seems more probable that they have passed through this stage, as during the early stages of the transformation, the other parts of the cell, with the included zooid, could hardly have disappeared at once. In many cases the vibracula have a grooved support at the base, which seems to represent the fixed beak; though this support in some species is quite absent. This view of the development of the vibracula, if trustworthy, is interesting; for supposing that all the species provided with avicularia had become extinct, no one with the most vivid imagination would ever have thought that the vibracula had originally existed as part of an organ, resembling a bird’s head, or an irregular box or hood. It is interesting to see two such widely different organs developed from a common origin; and as the movable lip of the cell serves as a protection to the zooid, there is no difficulty in believing that all the gradations, by which the lip became converted first into the lower mandible of an avicularium, and then into an elongated bristle, likewise served as a protection in different ways and under different circumstances.

In the vegetable kingdom Mr. Mivart only alludes to two cases, namely the structure of the flowers of orchids, and the movements of climbing plants. With respect to the former, he says: “The explanation of their _origin_ is deemed thoroughly unsatisfactory—utterly insufficient to explain the incipient, infinitesimal beginnings of structures which are of utility only when they are considerably developed.” As I have fully treated this subject in another work, I will here give only a few details on one alone of the most striking peculiarities of the flowers of orchids, namely, their pollinia. A pollinium, when highly developed, consists of a mass of pollen-grains, affixed to an elastic foot-stalk or caudicle, and this to a little mass of extremely viscid matter. The pollinia are by this means transported by insects from one flower to the stigma of another. In some orchids there is no caudicle to the pollen-masses, and the grains are merely tied together by fine threads; but as these are not confined to orchids, they need not here be considered; yet I may mention that at the base of the orchidaceous series, in Cypripedium, we can see how the threads were probably first developed. In other orchids the threads cohere at one end of the pollen-masses; and this forms the first or nascent trace of a caudicle.

That this is the origin of the caudicle, even when of considerable length and highly developed, we have good evidence in the aborted pollen-grains which can sometimes be detected embedded within the central and solid parts.

With respect to the second chief peculiarity, namely, the little mass of viscid matter attached to the end of the caudicle, a long series of gradations can be specified, each of plain service to the plant. In most flowers belonging to other orders the stigma secretes a little viscid matter. Now, in certain orchids similar viscid matter is secreted, but in much larger quantities by one alone of the three stigmas; and this stigma, perhaps in consequence of the copious secretion, is rendered sterile. When an insect visits a flower of this kind, it rubs off some of the viscid matter, and thus at the same time drags away some of the pollen-grains. From this simple condition, which differs but little from that of a multitude of common flowers, there are endless gradations—to species in which the pollen-mass terminates in a very short, free caudicle—to others in which the caudicle becomes firmly attached to the viscid matter, with the sterile stigma itself much modified. In this latter case we have a pollinium in its most highly developed and perfect condition. He who will carefully examine the flowers of orchids for himself will not deny the existence of the above series of gradations—from a mass of pollen-grains merely tied together by threads, with the stigma differing but little from that of the ordinary flowers, to a highly complex pollinium, admirably adapted for transportal by insects; nor will he deny that all the gradations in the several species are admirably adapted in relation to the general structure of each flower for its fertilisation by different insects. In this, and in almost every other case, the enquiry may be pushed further backwards; and it may be asked how did the stigma of an ordinary flower become viscid, but as we do not know the full history of any one group of beings, it is as useless to ask, as it is hopeless to attempt answering, such questions.

We will now turn to climbing plants. These can be arranged in a long series, from those which simply twine round a support, to those which I have called leaf-climbers, and to those provided with tendrils. In these two latter classes the stems have generally, but not always, lost the power of twining, though they retain the power of revolving, which the tendrils likewise possess. The gradations from leaf-climbers to tendril bearers are wonderfully close, and certain plants may be differently placed in either class. But in ascending the series from simple twiners to leaf-climbers, an important quality is added, namely sensitiveness to a touch, by which means the foot-stalks of the leaves or flowers, or these modified and converted into tendrils, are excited to bend round and clasp the touching object. He who will read my memoir on these plants will, I think, admit that all the many gradations in function and structure between simple twiners and tendril-bearers are in each case beneficial in a high degree to the species. For instance, it is clearly a great advantage to a twining plant to become a leaf-climber; and it is probable that every twiner which possessed leaves with long foot-stalks would have been developed into a leaf-climber, if the foot-stalks had possessed in any slight degree the requisite sensitiveness to a touch.

As twining is the simplest means of ascending a support, and forms the basis of our series, it may naturally be asked how did plants acquire this power in an incipient degree, afterwards to be improved and increased through natural selection. The power of twining depends, firstly, on the stems while young being extremely flexible (but this is a character common to many plants which are not climbers); and, secondly, on their continually bending to all points of the compass, one after the other in succession, in the same order. By this movement the stems are inclined to all sides, and are made to move round and round. As soon as the lower part of a stem strikes against any object and is stopped, the upper part still goes on bending and revolving, and thus necessarily twines round and up the support. The revolving movement ceases after the early growth of each shoot. As in many widely separated families of plants, single species and single genera possess the power of revolving, and have thus become twiners, they must have independently acquired it, and cannot have inherited it from a common progenitor. Hence, I was led to predict that some slight tendency to a movement of this kind would be found to be far from uncommon with plants which did not climb; and that this had afforded the basis for natural selection to work on and improve. When I made this prediction, I knew of only one imperfect case, namely, of the young flower-peduncles of a Maurandia which revolved slightly and irregularly, like the stems of twining plants, but without making any use of this habit. Soon afterwards Fritz Müller discovered that the young stems of an Alisma and of a Linum—plants which do not climb and are widely separated in the natural system—revolved plainly, though irregularly, and he states that he has reason to suspect that this occurs with some other plants. These slight movements appear to be of no service to the plants in question; anyhow, they are not of the least use in the way of climbing, which is the point that concerns us.

Nevertheless we can see that if the stems of these plants had been flexible, and if under the conditions to which they are exposed it had profited them to ascend to a height, then the habit of slightly and irregularly revolving might have been increased and utilised through natural selection, until they had become converted into well-developed twining species.

With respect to the sensitiveness of the foot-stalks of the leaves and flowers, and of tendrils, nearly the same remarks are applicable as in the case of the revolving movements of twining plants. As a vast number of species, belonging to widely distinct groups, are endowed with this kind of sensitiveness, it ought to be found in a nascent condition in many plants which have not become climbers. This is the case: I observed that the young flower-peduncles of the above Maurandia curved themselves a little towards the side which was touched. Morren found in several species of Oxalis that the leaves and their foot-stalks moved, especially after exposure to a hot sun, when they were gently and repeatedly touched, or when the plant was shaken. I repeated these observations on some other species of Oxalis with the same result; in some of them the movement was distinct, but was best seen in the young leaves; in others it was extremely slight. It is a more important fact that according to the high authority of Hofmeister, the young shoots and leaves of all plants move after being shaken; and with climbing plants it is, as we know, only during the early stages of growth that the foot-stalks and tendrils are sensitive.

It is scarcely possible that the above slight movements, due to a touch or shake, in the young and growing organs of plants, can be of any functional importance to them. But plants possess, in obedience to various stimuli, powers of movement, which are of manifest importance to them; for instance, towards and more rarely from the light—in opposition to, and more rarely in the direction of, the attraction of gravity. When the nerves and muscles of an animal are excited by