Behavior of the Lower Organisms

2. Interference with these processes causes a change of behavior and varied movements, subjecting the organism to many different conditions. 3. One of these conditions relieves the interference with the internal processes, so that the changes in behavior cease. It is clear that regulation taking place in this way does not require that the end or purpose of the action shall function in any way as part of its cause, as is held in various vitalistic theories. There is no evidence that a final aim is guiding the organism. None of the factors above mentioned appear to include anything differing in essential principle from such methods of action as we find in the inorganic world.

Now an additional factor enters the problem. By the process which we have just considered, the organism reaches in time a movement that brings relief from the interfering conditions. This relieving response becomes fixed through the operation of the law of the readier resolution of physiological states as a result of repetition (Chapter XVI, Section 10). After reaching the relieving response a number of times by a repeated succession of movements, a recurrence of the interfering condition induces more quickly the relieving response, and in time this becomes the immediate reaction to this interfering condition.

It is in this second stage of the process, when the relieving response has become set through the law of the readier resolution of physiological states by repetition, that an end or purpose seems to dominate the behavior. This end or purpose of course actually exists, as a subjective state called an idea, in man. Whether any such subjective state exists in the lower organism that has gone through the process just sketched, of course we do not know. But some objective phenomenon, as a transient physiological state, would seem to be required in the lower animal, corresponding to the objective physiological accompaniment of the idea in man. The behavior in this stage is that which, in its higher reaches at least, has been called intelligent.

But so far as the objective occurrences are concerned, there would seem to be nothing in this later stage of the behavior involving anything different in essential principle from what we find in the inorganic world. The only additional factor is the law of the readier resolution of physiological states after repetition. While possibly our statement of this law may not be entirely adequate, there would seem to be nothing implied by it that is specifically vital, in the sense that it differs in essential principle from the methods of action seen in the inorganic world. This law of the readier resolution of physiological states after repetition presents indeed many analogies with various chains of physical and chemical action.1 It certainly by no means requires in itself the action of any "final cause," — that is, of an entity that is at the same time purpose and cause. On the other hand, it undoubtedly does produce that type of behavior which has given rise to the conception of the purpose acting as cause. This conception is in itself of course a correct one, so far as we mean by a purpose an actual physiological state of the organism, determining behavior in the same manner as other factors determine it. But such a physiological state (subjectively a purpose) is a result of a foregoing objective cause, and acts to produce an effect in the same way as any other link in the causal chain. It would seem therefore to present no basis for theories of vitalism, so far as these depend on anything like the action of final causes.

That regulation takes place in the behavior of many animals in the manner above sketched may be affirmed as a clearly established fact, and it seems to be perhaps the only intelligible way in which regulatory behavior could be developed in a given individual. But we are, of course, confronted by the fact that many individuals are provided at birth with definite regulatory methods of reaction to certain stimuli. In these cases the animal is not compelled to go through the process of performing varied movements, with subsequent fixation of the successful movement. How are such cases to be accounted for?

If the regulatory method of reaction acquire! through the process sketched in the preceding paragraphs could be inherited, there would of course be no difficulty in accounting for such congenital regulatory reactions. In Protozoa this is apparently the real state of the case ; there appears to be no reason why the products of reproduction by division should not inherit the properties of the individual that divides, however these properties were attained. But in the Metazoa such inheritance of acquirements presents great theoretical difficulties, and has not been experimentally demonstrated to occur, though it is perhaps too early to consider the matter as yet out of court. If such inheritance does not occur, the existence of congenital definite regulatory reactions would seem explicable only on the basis of the natural selection of individuals having varying methods of reaction, unless we are to adopt the theories of vitalism. In the method we have sketched above, a certain reaction that is

regulatory is selected, through the operation of physiological laws, from among many performed by the same individual. In natural selection the same reaction is selected from among many performed by different individuals — in both cases because it is regulatory — because it assists the life processes of the organism. The two factors must then work together and produce similar results.1 In both, the essential point is a selection from among varied activities.

We must here notice the fact that we often find in organisms behavior that is not regulatory. How are we to account for this ? Without going into details, it is clear that there are a number of factors that would produce this result. First, interference with the life processes is not the only cause of reaction. The organism is composed of matter that is subject to the usual laws of physics and chemistry. External agents may of course act on this matter directly, causing changes in movement that are not regulatory. Second, the organism can perform only those movements which its structure permits. Often none of these movements can produce conditions that relieve the existing interference with the life processes. Then the organism can only try them, without regulatory results, and die (see, for example, such a case in the flatworm, p. 244). Further, certain responses may have become fixed, in the way described above, because under usual conditions they produce adjustment. Now if the conditions change, the organism still responds by the fixed reaction, and this may no longer be regulatory. The organism may then be destroyed before a new regulatory reaction can be developed by selection from varied movements. This condition of affairs is of course often observed.

All together, the regulatory character of behavior as found in many animals seems intelligible in a perfectly natural, directly causal way, on the basis of the principles brought out above. We may summarize these principles as (1) the selection through varied movements of conditions not interfering with the physiological processes of the organism ("trial and error"); (2) the fixation of the adaptive movements through the law of the readier resolution of physiological states after repetition.

Is it possible that individual regulation in other fields is based on the same principles that we have set forth above for behavior? Bodily movement is only one of the many activities that vary, and variations of any of the organic activities may impede or assist the physiological 1 For a discussion of the relation of these two factors, see Chapter XIX. processes of the organism. Is it possible that interference with the physiological processes may induce changes in other activities, — in chemical processes, in growth, and the like, — and that one of these activities is selected, as in behavior, through the fact that it relieves the interference that caused the change ?

There is some evidence for this possibility. Let us look, for example, at regulative changes in the chemical activity of the organism, such as we see in the acclimatization to poisons, in the responses to changes in temperature, or in the adaptation of the digestive juices to the food. What is the material from which the regulative conditions may be selected? One of the general results of modern physical chemistry is expressed by Ostwald (1902, p. 366) as follows: "In a given chemical structure all processes that are so much as possible, are really taking place, and they lead to the formation of all substances that can occur at all." Some of these processes are taking place so slowly that they escape usual observation ; we notice only those that are conspicuous. But in its enzymes the body possesses the means (as Ostwald sets forth) of hastening any of these processes and delaying others, so that the general character of the action shall be determined by the more rapid process. Such enzymes are usually present in the body in inactive forms (zymogens), which may be transformed into active enzymes by slight chemical changes, thus altering fundamentally the course of the chemical processes in the organisms.

It is evident, then, that the organism has presented to it, by the condition just sketched, unlimited possibilities for the selection of different chemical processes. The body is a great mass of the most varied chemicals, and in this mass thousands of chemical processes, in every direction, — all those indeed that are possible, — are occurring at all times. There is then no difficulty as to the sufficiency of the material presented for selection, if some means may be found for selecting it.

Further, it is known that interference with the physiological processes does result in many changes in the internal activities of the organism, as well as in its external movements. Intense injurious stimulation causes not merely excess movements of the body as a whole, but induces marked changes in circulation, in respiration, in temperature, in digestive processes, in excretion, and in other ways. Such marked internal changes involve, and indeed are constituted by, alterations of profound character in the chemical processes of the organism. These chemical changes are sometimes demonstrated by the production of new chemicals under such circumstances. Furthermore, it is clear that the internal changes due to interference with the physiological processes are not stereotyped in character, but varied. Under violent injurious stimu-

lation, respiration becomes for a time rapid, then is almost suspended. The heart beats for a time furiously, then feebly, and there is similar variation in other internal symptoms. Thus it seems clear that interference with the life processes does produce varied activities in other ways than in bodily movements ; and that among these it results in varied chemical processes. There is then presented opportunity for regulation to occur in the same way as in behavior. Certain of the processes occurring relieve the disturbance of the physiological functions. There results a cessation of the changes. In other words, a certain process is selected through the fact that it does relieve. It is well known, through the work of Pawlow (1898), that the adaptive changes in the activities of the digestive glands, fitting the digestive juices to the food taken, do not occur at once and completely under a given diet, but are brought about gradually. As the dog is continued on a diet of bread, the pancreatic juice becomes more and more adapted to the digestion of starch. This slow adaptation is of course what should be expected if the process occurs in anything like the manner we have sketched.

At a later stage, if the laws of these processes are the same as those for behavior, there will be present certain fixed methods of chemical response, by which the organism reacts to certain sorts of stimulation. That the law of the readier resolution of physiological states after repetition holds in this field, is clearly indicated by the work of Pawlow. He found that the pancreas under a uniform diet does tend to acquire a fixed method of reaction to the introduction of the food, that is not easily changed. In the dog which has digested starch for a month, the pancreatic juice is not readily changed back to that adapted to the digestion of meat. As a result, definite organs will in the course of time have left open to them only certain limited possibilities of variation — due to the development of something corresponding to the "action system" in behavior. Thus, in the pancreas, there will not exist unlimited possibilities as to the chemical changes that may occur. Its "action system" will be limited perhaps to the production of varied quantities of a certain set of enzymes, — amylopsin, trypsin, etc. The proper selection of these few possibilities will then occur by the method sketched. When digestion is disturbed by food that is not well digested, variations in the production of the different enzymes will be set in train, and one of these will in time relieve the difficulty, through the more complete digestion of the food. Thereupon the variations will cease, since their cause has disappeared. By still more complete fixation of the chemical response, through the law of the readier resolution of physiological states after repetition, or the analogue of this law, an organ or organism may

largely lose its power of varying its chemical behavior and thus be unable to meet new conditions in a regulative way. A condition comparable to the production of a fixed reflex in behavior will result. It is perhaps more difficult to apply the method of regulation above set forth to processes of growth and regeneration. Yet there is no logical difficulty in the way. The only question would be that of fact, — whether the varied growth processes necessary do, primitively, occur under conditions that interfere with the physiological processes. When a wound is made or an organ removed, is the growth process which follows always of a certain stereotyped character, or are there variations ? It. is well known, of course, that the latter is the case. In the regeneration of the earthworm, Morgan (1897) finds great variation; he says that in trying many experiments, one finds that what ninety-nine worms cannot do in the way of regeneration, the one hundredth can. The very great variations in the results of operations on eggs and young stages of animals are well known. Removal of an organ is known to produce great disturbance of most of the processes in the organism, and among others in the process of growth.

It appears not impossible then that regulation may be brought about in growth processes in accordance with the same principles as in behavior. A disturbance of the physiological processes results in varied activities, and among these are varied growth activities. Some of these relieve the disturbance; the variation then ceases and these processes are continued. In any given highly organized animal or plant the different possibilities of growth will have become decidedly limited ; and it is only from this limited number of possibilities that selections can be made. In some cases, by the fixation of certain processes through the analogue of the law of the readier resolution of physiological states, the organism or a certain part thereof will have lost the power of responding to injury save in one definite way. Under new conditions this one way may not be regulatory, yet it may be the only response possible. Thus may result the formation under certain conditions of heteromorphic structures, — a tail in place of a head, or the like, from a part of the body that (in normal development perhaps) is accustomed to produce such an organ. This would again correspond to the production of a fixed reflex action in behavior, even under circumstances where this action is not regulatory.

It appears to the writer that the method of form regulation recently set forth in a most suggestive paper by Holmes (1904) is in agreement with the general method of regulation here set forth, and may be considered a working out of the details of the way in which growth regulation might take place along these lines. Holmes has of course emphasized other features of the process in a way that is not called for in the present work. Some suggestions as to the possibility of regulation along the line of the selection of overproduced activities are found in J. Mark Baldwin's valuable collection of essays entitled "Evolution and Development."

It may be noted that regulation in the manner we have set forth is what in behavior is commonly called intelligence. If the same method of regulation is found in other fields, then there is no reason for refusing to compare the action there to intelligence. Comparison of the regulatory processes that are shown in internal physiological changes and in regeneration, to intelligence seems to be looked upon sometimes as unscientific and heretical. Yet intelligence is a name applied to processes that actually exist in the regulation of movement, and there is no a priori reason why similar processes should not occur in regulation in other fields. Movement is after all only the general result of the more recondite chemical and physical changes occurring in organisms, and therefore cannot follow laws differing in essential character from the latter. We are dealing in other fields with the same substance that is capable of performing the processes seen in intelligent action, and these could not occur as they do if the underlying physical and chemical processes did not obey the same laws. In a purely objective consideration there seems no reason to suppose that regulation in behavior (intelligence) is of a fundamentally different character from regulation elsewhere.

We may sum up the fundamental features in the method of individual regulation above set forth as follows : — The organism is a complex of many processes, of chemical change, of growth, and of movement ; these are proceeding with a certain energy. These processes depend for their unimpeded course on their relations to each other and on the relations to the environment which the processes themselves bring about. When any of these processes are blocked or disturbed, through a change in the relations to each other or the environment, the energy overflows in other directions, producing varied changes, — in movement, and apparently also in chemical and growth processes. These changes of course vary the relations of the processes to each other and to the environment; some of the conditions thus reached relieve the interference which was the cause of the change. Thereupon the changes cease, since there is no further cause for them; the relieving condition is therefore maintained. After repetition of this course of events, the process which leads to relief is reached more

directly, as a result of the law of the readier resolution of physiological states after repetition. Thus are produced finally the stereotyped changes often resulting from stimulation. This method of regulation is clearly seen in behavior, where its operation is, in the later stages, what is called intelligence. Its application to chemical and form regulation is at present hypothetical, but appears possible. The following is a list of the works cited in the text. It is not a complete bibliography of behavior in lower animals, but will be found to contain most of the more important papers on the lowest groups. The authors' names are given in alphabetical order, with their works arranged according to the date of their appearance. In the text the works are cited by the name of the author accompanied by the date; for the complete title reference is to be made to the present list.

Adams, G. P., 1903. On the negative and positive phototropism of the earthworm Allolobophora fcetida (Sav.), as determined by light of different intensities: Amer. Journ. Physiol., IX, 26-34. — Allabach, L. F., 1905. Some points regarding the behavior of Metridium: Biol. Bui., X, 35-43. Antholoba reticulata Couth, und Hepatus chilensis M. E. : Biol. Centralbl., XXIII, 677-678. — Butschli, O., 1880. Protozoa, I Abth. Bronn's Klassen und Ordnungen des Thierreichs. Leipzig. — Id., 1889. Protozoa, III Abth. Infusoria. Bronn's Klassen und Ordnungen des Thierreichs, I Bd. Leipzig. — Id., 1892. Untersuchungen iiber mikroskopische Schaume und das Protoplasma. 234 pp. Leipzig.

Carlgren, O., 1899. Ueber die Eimvirkung des constanten galvanischen Stromes auf niedere Organismen: Arch. f. Anat. u. Physiol. Physiol. Abth., 49-76. — Id., 1905. Ueber die Bedeutung der Flimmerbewegung fiir die Xahrungstransport bei den Actiniarien und Madreporarien : Biol. Centralbl., XXV, 308-322. — Id., 1905 a. Der Galvanotropismus und die innere Kataphorese: Zeitschr. f. allg. Physiol., V, 123-130. — Claparede, Ed., 1901. Les animaux sont-ils conscients?: Revue Philosophique, LI, 24 pp. (Translation in International Quarterly, VIII, 296-315.) — Id., 1905. La psychologie comparee est-elle legitime?: Archives d. Psychol., V, 13-35. — Coehn, A. and Barratt, W., 1905. Ueber Galvanotaxis vora Standpunkte der physikalische Chemie: Zeitschr. f. allg. Physiol., V, 1-9.

Famintzin, A., 1867. Die Wirkung des Lichtes auf Algen und einige andere nahe verwandte Organismen: Jahrb. f. wiss. Bot., VI, 1-44. Gamble, F. W. and Keeble, F., 1903. The bionomics of Convoluta roscoffensis, with special reference to its green cells: Quart. Journ. Micr. Sci., XL VII, 363- 431. — Garrey, W. E., 1900. The effect of ions upon the aggregation of flagellated infusoria: Amer. Journ. Physiol., Ill, 291-315. — Greeley, A. W., 1904. Experiments on the physical structure of the protoplasm of Paramecium and its relation to the reactions of the organism to thermal, chemical, and electrical stimuli : Biol. Bui. VII, 3-32.

Haberlandt, G., 1905. Ueber den Begriff "Sinncsorgane" in der Tierund Pflanzenphysiologie : Biol. Centralbl., XXV, 446-451. — Harper, E. EL, 1905. Reactions to light and mechanical stimuli in the earthworm, Perichaeta bermudensis (Beddard): Biol. Bui., X, 17-34. — Harrington, Xt. R. and Leaming, E., 1900. The reaction of Amoeba to light of different colors: Amer. Journ. Physiol., Ill, 9-16. — Hertel, E., 1904. Ueber Beeinflussung des Organismus durch Licht,

Kuhne, W., 1864. Untersuchungen liber das Protoplasma und die Contractilitait. 158 pp. Leipzig. Le Dantec, F., 1895. La matiere vivante. 191 pp. Paris. — Leidy, J., 1879. Fresh -water rhizopods of North America: Rep. U. S. Geol. Survey of the Territories, XII, 334 pp., 48 pi. — Loeb, J., 1891. Untersuchungen zur physiologischen Morphologie der Thiere. I. Ueber Heteromorphose. 80 pp. Wiirzburg. — Id., 1893. Ueber klinstliche Umwandlung positiv heliotropischer Thiere in negativ heliotropische und umgekehrt: Arch. f. d. ges. Physiol., LIV, 81-107. — ID-> 1894. Beitrage zur Gehirn physiologie der Wiirmer: Arch. f. d. ges. Physiol., LVI, 247-269. — Id., 1895. Zur Physiologie und Psychologie der Actinien : Arch. f. d. ges. Physiol., LIX, 415-420. — Id., 1897. Zur Theorie der physiologischen Licht und Schwerkraftwirkungen : Arch. f. d. ges. Physiol., LXVI, 439-466. — Id., 1900. Comparative physiology of the brain and comparative psychology. 309 pp. New York. — Id., 1900 a. On the different effects of ions upon myogenic and neurogenic rhythmical contractions and upon embryonic and muscular tissue: Amer. Journ. Physiol., Ill, 384-396. — ■ Loeb, J. and Budgett, S. P., 1897. Zur Theorie der Galvanotropismus. IV. Mittheilung. Ueber die Ausscheidung electropositiver Ionen an den ausseren Anodenflache protoplasmatischer Gebilde als Ursache der Abweichungen vom Pflliger'schen Erregungsgesetz: Arch. f. d. ges. Physiol., LXVI, 518-534. — Ludloff, K., 1895. Untersuchungen liber den Galvanotropismus: Arch. f. d. ges. Physiol., LIX, 525-554. — Ltjkas, F., 1905. Psychologie der niedersten Tiere. 276 pp. Wien und Leipzig. — Lyon, E. P., 1904. On rheotropism. I. Rheotropism in fishes: Amer. Journ. Physiol., XII, 149-161. — Id., 1905. On the theory of geotropism in Paramcecium: Amer. Journ. Physiol., XIV,

Marshall, W., 1882. Ueber einige Lebenserscheinungen der Siisswasserpolypen und liber eine neue Form von Hydra viridis : Zeitschr. f. wiss. Zool., XXXVII, 664-702. — Massart, J., 1889. Sensibilite et adaptation des organismes a la concentration des solutions salines: Arch, de Biol., IX, 515-570. — Id., 1891. Recherches sur les organismes inferieurs. II. La sensibilite a la concentration chez les etres unicellulaires marins: Bui. Acad. roy. Sci. Belgique, (3), XXII, 158- 167. — Id., 1891 a. Recherches sur les organismes inferieurs. III. La sensibilite a la gravitation: Bui. Acad. roy. Belgique (3), XXII, 158-167. — Id., 1901. Essai

Naegeli, C, i860. Ortsbewegungen der Pflanzenzellen und ihrer Theile (Stromungen) : Naegeli's Beitrage zur wiss. Bot, Hft. II, 59-108. — Nagel, W., 1892. Das Geschmacksinn der Actinien: Zool. Anz., XV, 334-338- — Id., 1894. Vergleichendphysiologische und anatomische Untersuchungen iiber den Geruchsund Geschmacksinn und ihre Organe: Bibliotheca Zoologica, XVIII, 207 pp. — Id., 1894 a. Experimented sinnesphysiologische Untersuchungen an Coelenteraten : Arch. f. d. ges. Physiol., LVII, 495-552. — Id., 1899. Ueber neue Nomenclatur in der vergleichenden Sinnesphysiologie : Centralbl. f. Physiol., Hft. XII., 4 pp. — Nuel, 1904. La vision. 376 pp. Paris.

Radl, E., 1903. Untersuchungen iiber die Phototropismus der Thiere. 188 pp. Leipzig. — Rhumbler, L., 1898. Physikalische Analyse von Lebenserscheinungen der Zelle I. Bewegung, Nahrungsaufnahme, Defalcation, Vacuolen-Pulsationen, und Gehausebau bei lobosen Rhizopoden: Arch. f. Entw.-mech., VII, 103-350.— Id., 1905. Zur Theorie der Oberflachenkrafte der Amdben: Zeitschr. f. wiss Zool., LXXXIII, 1-52. — Roesle, E., 1902. Die Reaktion einiger Infusorien auf einzelne Induktionsschlage : Zeitschr. f. allg. Physiol, II, 139-168. — Romanes, G. J., 1885. Jellyfish, starfish, and sea urchins. 323 pp. New York. — Rothert, W., 1901. Beobachtungen und Betrachtungen iiber tactische Reizerscheinungen : Flora, LXXXVIII, 371-421. — Id., 1903. Ueber die Wirkung des Aethers und Chloroforms auf die Reizbewegungen der Mikroorganismen : Jahrb. f. wiss. Bot., XXXIX, 1-70. — Roux, W., 1901. Ueber die " morphologische Polarisation" von Eiern und Em- bryonen durch den electrischen Strom: Sitz.-ber. d. k. Akad. d. Wiss. z. Wien, Math, u. Naturw. Classe, CI, 27-228. (Roux's Gesammelte Abhandlungen, II, 540-765.)

Schwarz, F., 1884. Der Einfluss der Schwerkraft auf die Bewegungsrichtung von Chlamidomonas und Euglena: Ber. Bot. Gesellsch., II, 51-72. — Semon, R., 1904. Die Mneme als erhaltendes Prinzip im Wechsel des organischen Geschehens. 353 pp. Leipzig. — Smith, Amelia C, 1902. The influence of temperature, odors, light, and contact on the movements of the earthworm: Amer. Journ. Physiol., VI, 459-486. — SosNOWSKi, J., 1899. Untersuchungen iiber die Verainderungen der Geotropismus bei Paramecium aurelia: Bui. Internat. Acad. Sci. Cracovie, 130-136. — Spaulding, E. G., 1904. An establishment of association in hermit crabs, Eupagurus longicarpus: Journ. Comp. Neurol, and Psychol., XIV, 49-61. — Spencer, H., 1894. Principles of psychology, 3d ed. 2 vols. New York. — Stahl, E., 1884. Zur Biologie der Myxomyceten: Bot. Zeitung, XL, 146-155; 162-175; 187-191. — Statkewitsch, P., 1903. Ueber die Wirkung der Induktionsschlage auf einige Ciliata: Le Physiologiste Russe, III, 55 pp. — Id., 1903 a. Galvanotropism and galvanotaxis of organisms. Part First. Galvanotropism and