FOOTNOTES:[3]As Whitman has pointed out, Aristotle taught epigenesis as clearly as Harvey, and is, therefore, to be regarded as the founder of that conception.[4]The discovery is also attributed to Hamm, a medical student, and to Hartsoeker, who claimed priority in the discovery.[5]De Formatione Intestinorum, Nova Commentar, Ac. Sci. Petrop., St. Petersburg, XII., 1768; XIII., 1769.[6]Besides biographical sketches by Stieda, Waldeyer, and others, we have a very entertaining autobiography of Von Baer, published in 1864, for private circulation, but afterward (1866) reprinted and placed on sale.[7]It is of more than passing interest to remember that Pander and Von Baer were associated as friends and fellow-students, under Döllinger at Würzburg. It was partly through the influence of Von Baer that Pander came to study with Döllinger, and took up investigations on development. His ample private means made it possible for him to bear the expenses connected with the investigation, and to secure the services of a fine artist for making the illustrations. The result was a magnificently illustrated treatise. His unillustrated thesis in Latin (1817) is more commonly known, but the illustrated treatise in German is rarer. Von Baer did not take up his researches seriously until Pander's were published. It is significant of their continued harmonious relations that Von Baer's work is dedicated "An meinen Jugendfreund, Dr. Christian Pander."
FOOTNOTES:
[3]As Whitman has pointed out, Aristotle taught epigenesis as clearly as Harvey, and is, therefore, to be regarded as the founder of that conception.
[3]As Whitman has pointed out, Aristotle taught epigenesis as clearly as Harvey, and is, therefore, to be regarded as the founder of that conception.
[4]The discovery is also attributed to Hamm, a medical student, and to Hartsoeker, who claimed priority in the discovery.
[4]The discovery is also attributed to Hamm, a medical student, and to Hartsoeker, who claimed priority in the discovery.
[5]De Formatione Intestinorum, Nova Commentar, Ac. Sci. Petrop., St. Petersburg, XII., 1768; XIII., 1769.
[5]De Formatione Intestinorum, Nova Commentar, Ac. Sci. Petrop., St. Petersburg, XII., 1768; XIII., 1769.
[6]Besides biographical sketches by Stieda, Waldeyer, and others, we have a very entertaining autobiography of Von Baer, published in 1864, for private circulation, but afterward (1866) reprinted and placed on sale.
[6]Besides biographical sketches by Stieda, Waldeyer, and others, we have a very entertaining autobiography of Von Baer, published in 1864, for private circulation, but afterward (1866) reprinted and placed on sale.
[7]It is of more than passing interest to remember that Pander and Von Baer were associated as friends and fellow-students, under Döllinger at Würzburg. It was partly through the influence of Von Baer that Pander came to study with Döllinger, and took up investigations on development. His ample private means made it possible for him to bear the expenses connected with the investigation, and to secure the services of a fine artist for making the illustrations. The result was a magnificently illustrated treatise. His unillustrated thesis in Latin (1817) is more commonly known, but the illustrated treatise in German is rarer. Von Baer did not take up his researches seriously until Pander's were published. It is significant of their continued harmonious relations that Von Baer's work is dedicated "An meinen Jugendfreund, Dr. Christian Pander."
[7]It is of more than passing interest to remember that Pander and Von Baer were associated as friends and fellow-students, under Döllinger at Würzburg. It was partly through the influence of Von Baer that Pander came to study with Döllinger, and took up investigations on development. His ample private means made it possible for him to bear the expenses connected with the investigation, and to secure the services of a fine artist for making the illustrations. The result was a magnificently illustrated treatise. His unillustrated thesis in Latin (1817) is more commonly known, but the illustrated treatise in German is rarer. Von Baer did not take up his researches seriously until Pander's were published. It is significant of their continued harmonious relations that Von Baer's work is dedicated "An meinen Jugendfreund, Dr. Christian Pander."
CHAPTER XI
THE CELL THEORY—SCHLEIDEN, SCHWANN, SCHULTZE
Therecognition, in 1838, of the fact that all the various tissues of animals and plants are constructed on a similar plan was an important step in the rise of biology. It was progress along the line of microscopical observation. One can readily understand that the structural analysis of organisms could not be completed until their elementary parts had been discovered. When these units of structure were discovered they were called cells—from a misconception of their nature—and, although the misconception has long since been corrected, they still retain this historical but misleading name.
The doctrine that all tissues of animals and plants are composed of aggregations of these units, and the derivatives from the same, is known as the cell-theory. It is a generalization which unites all animals and plants on the broad plane of similitude of structure, and, when we consider it in the light of its consequences, it stands out as one of the great scientific achievements of the nineteenth century. There is little danger of overestimating the importance of this doctrine as tending to unify the knowledge of living organisms.
Vague Foreshadowings of the Cell-Theory.—In attempting to trace the growth of this idea, as based on actual observations, we first encounter vague foreshadowings of it in the seventeenth and the eighteenth centuries. The cells were seen and sketched by many early observers, but were not understood.
As long ago as 1665 Robert Hooke, the great English microscopist, observed the cellular construction of cork, and described it as made up of "little boxes or cells distinguished from one another." He made sketches of the appearance of this plant tissue; and, inasmuch as the drawings of Hooke are the earliest ones made of cells, they possess especial interest and consequently are reproduced here. Fig. 72, taken from theMicrographia, shows this earliest drawing of Hooke. He made thin sections with a sharp penknife; "and upon examination they were found to be all cellular or porous in the manner of a honeycomb, but not so regular."
Fig. 72.—The Earliest Known Picture of Cells from Hooke'sMicrographia(1665). From the edition of 1780.
We must not completely overlook the fact that Aristotle (384-322B.C.) and Galen (130-200A.D.), those profound thinkers on anatomical structure, had reached the theoretical position "that animals and plants, complex as they mayappear, are yet composed of comparatively few elementary parts, frequently repeated"; but we are not especially concerned with the remote history of the idea, so much as with the principal steps in its development after the beginning of microscopical observations.
Fig. 73.—Sketch from Malpighi's Treatise on the Anatomy of Plants (1670).
Pictures of Cells in the Seventeenth Century.—The sketches illustrating the microscopic observations of Malpighi, Leeuwenhoek, and Grew show so many pictures of the cellular construction of plants that one who views them for the first time is struck with surprise, and might readily exclaim: "Here in the seventeenth century we have the foundation of the cell-theory." But these drawings were merely faithful representations of the appearance of the fabric of plants;the cells were not thought of as uniform elements of organic architecture, and no theory resulted. It is true that Malpighi understood that the cells were separable "utricles," and that plant tissue was the result of their union, but this was only an initial step in the direction of the cell-theory, which, as we shall see later, was founded on the supposed identity in development of cells in animals and plants. Fig. 73 shows a sketch, made by Malpighi about 1670, illustrating the microscopic structure of a plant. This is similar to the many drawings of Grew and Leeuwenhoek illustrating the structure of plant tissues.
Wolff.—Nearly a century after the work of Malpighi, we find Wolff, in 1759, proposing a theory regarding the organization of animals and plants based upon observations of their mode of development. He was one of the most acute scientific observers of the period, and it is to be noted that his conclusions regarding structure were all founded upon what he was able to see; while he gives some theoretical conclusions of a purely speculative nature, Wolff was careful to keep these separate from his observations. The purpose of his investigations was to show that there was no pre-formation in the embryo; but in getting at the basis of this question, he worked out the identity of structure of plants and animals as shown by their development. In his famous publication on the Theory of Development (Theoria Generationis) he used both plants and animals.
Huxley epitomizes Wolff's views on the development of elementary parts as follows: "Every organ, he says, is composed at first of a little mass of clear, viscous, nutritive fluid, which possesses no organization of any kind, but is at most composed of globules. In this semifluid mass cavities (Bläschen,Zellen) are now developed; these, if they remain round or polygonal, become the subsequent cells; if they elongate, the vessels; and the process is identically the same,whether it is examined in the vegetating point of a plant, or in the young budding organs of an animal."
Wolff was contending against the doctrine of pre-formation in the embryo (see further under the chapter on Embryology), but on account of his acute analysis he should be regarded, perhaps, as the chief forerunner of the founders of the cell-theory. He contended for the same method of development that was afterward emphasized by Schleiden and Schwann. Through the opposition of the illustrious physiologist Haller his work remained unappreciated, and was finally forgotten, until it was revived again in 1812.
We can not show that Wolff's researches had any direct influence in leading Schleiden and Schwann to their announcement of the cell-theory. Nevertheless, it stands, intellectually, in the direct line of development of that idea, while the views of Haller upon the construction of organized beings are a side-issue. Haller declared that "the solid parts of animals and vegetables have this fabric in common, that their elements are either fibers or unorganized concrete." This formed the basis of the fiber-theory, which, on account of the great authority of Haller in physiology, occupied in the accumulating writings of anatomists a greater place than the views of Wolff.
Bichat, although he is recognized as the founder of histology, made no original observations on the microscopic units of the tissues. He described very minutely the membranes in the bodies of animals, but did not employ the microscope in his investigations.
Oken.—In the work of the dreamer Oken (1779-1851), the great representative of the German school of "Naturphilosophie," we find, about 1808, a very noteworthy statement to the effect that "animals and plants are throughout nothing else than manifoldly divided or repeated vesicles, as I shall prove anatomically at the proper time." This isapparently a concise statement of the cell-idea prior to Schleiden and Schwann; but we know that it was not founded on observation. Oken, as was his wont, gave rein to his imagination, and, on his part, the idea was entirely theoretical, and amounted to nothing more than a lucky guess.
Haller's fiber-theory gave place in the last part of the eighteenth century to the theory that animals and plants are composed of globules and formless material, and this globular theory was in force up to the time of the great generalization of Schleiden and Schwann. It was well expounded by Milne-Edwards in 1823, and now we can recognize that at least some of the globules which he described were the nucleated cells of later writers.
The Announcement of the Cell-Theory.—We are now approaching the time when the cell-theory was to be launched. During the first third of the nineteenth century there had accumulated a great mass of separate observations on the microscopic structure of both animals and plants. For several years botanists, in particular, had been observing and writing about cells, and interest in these structures was increasing. "We must clearly recognize the fact that for some time prior to 1838 the cell had come to be quite universally recognized as a constantly recurring element in vegetable and animal tissues, though little importance was attached to it as an element of organization, nor had its character been clearly determined" (Tyson).
Then, in 1838, came the "master-stroke in generalization" due to the combined labors of two friends, Schleiden and Schwann. But, although these two men are recognized as co-founders, they do not share honors equally; the work of Schwann was much more comprehensive, and it was he who first used the term cell-theory, and entered upon the theoretical considerations which placed the theory before the scientific world.
Schleiden was educated as a lawyer, and began the practice of that profession, but his taste for natural science was so pronounced that when he was twenty-seven years old he deserted law, and went back to the university to study medicine. After graduating in medicine, he devoted himself mainly to botany. He saw clearly that the greatest thing needed for the advancement of scientific botany was a study of plant organization from the standpoint of development. Accordingly he entered upon this work, and, in 1837, arrived at a new view regarding the origin of plant cells. It must be confessed that this new view was founded on erroneous observations and conclusions, but it was revolutionary, and served to provoke discussion and to awaken observation. This was a characteristic feature of Schleiden's influence upon botany. His work acted as a ferment in bringing about new activity.
The discovery of the nucleus in plant cells by Robert Brown in 1831 was an important preliminary step to the work of Schleiden, since the latter seized upon the nucleus as the starting-point of new cells. He changed the name of the nucleus to cytoblast, and supposed that the new cell started as a small clear bubble on one side of the nucleus, and by continued expansion grew into the cell, the nucleus, or cytoblast, becoming encased in the cell-wall. All this was shown by Nägeli and other botanists to be wrong; yet, curiously enough, it was through the help of these false observations that Schwann arrived at his general conclusions.
Schleiden was acquainted with Schwann, and in October, 1838, while the two were dining together, he told Schwann about his observations and theories. He mentioned in particular the nucleus and its relationship to the other parts of the cell. Schwann was immediately struck with the similarity between the observations of Schleiden and certain of his own uponanimaltissues. Together they went to hislaboratory and examined the sections of the dorsal cord, the particular structure upon which Schwann had been working. Schleiden at once recognized the nuclei in this structure as being similar to those which he had observed in plants, and thus aided Schwann to come to the conclusion that the elements in animal tissues were practically identical with those in plant tissues.
Schwann.—The personalities of the co-founders of the cell-theory are interesting. Schwann was a man of gentle, pacific disposition, who avoided all controversies aroused by his many scientific discoveries. In his portrait (Fig. 74) we see a man whose striking qualities are good-will and benignity. His friend Henle gives this description of him: "He was a man of stature below the medium, with a beardless face, an almost infantile and always smiling expression, smooth, dark-brown hair, wearing a fur-trimmed dressing-gown, living in a poorly lighted room on the second floor of a restaurant which was not even of the second class. He would pass whole days there without going out, with a few rare books around him, and numerous glass vessels, retorts, vials, and tubes, simple apparatus which he made himself. Or I go in imagination to the dark and fusty halls of the Anatomical Institute where we used to work till nightfall by the side of our excellent chief, Johann Müller. We took our dinner in the evening, after the English fashion, so that we might enjoy more of the advantages of daylight."
Schwann drew part of his stimulus from his great master, Johannes Müller. He was associated with him as a student, first in the University of Würzburg, where Müller, with rare discernment for recognizing genius, selected Schwann for especial favors and for close personal friendship. The influence of his long association with Müller, the greatest of all trainers of anatomists and physiologists of the nineteenth century, must have been very uplifting. A few years later,Schwann found himself at the University of Berlin, where Müller had been called, and he became an assistant in the master's laboratory. There he gained the powerful stimulus of constant association with a great personality.
Fig. 74.—Theodor Schwann, 1810-1882.
In 1839, just after the publication of his work on the cell-theory, Schwann was called to a professorship in the University of Louvain, and after remaining there nine years, was transferred to the University of Liège. He was highly respected in the university, and led a useful life, although after going to Belgium he published only one work—that on the uses of the bile. He was recognized as an adept experimenter and demonstrator, and "clearness, order, and method" are designated as the characteristic qualities of his teaching.
Fig. 75.—M. Schleiden, 1804-1881.
His announcement of the cell-theory was his most important work. Apart from that his best-known contributions to science are: experiments upon spontaneous generation, his discovery of the "sheath of Schwann," in nerve fibers, and his theory of fermentation as produced by microbes.
Schleiden.—Schleiden (Fig. 75) was quite different in temperament from Schwann. He did not have the fine self-control of Schwann, but was quick to take up the gauntlet and enter upon controversies. In his caustic replies to his critics, he indulged in sharp personalities, and one is at times inclined to suspect that his early experience as a lawyer had something to do with his method of handling opposition. With all this he had correct ideas of the object of scientific study and of the methods to be used in its pursuit. He insisted upon observation and experiment, and upon the necessity of studying the development of plants in order to understand their anatomy and physiology. He speaks scornfully of the botany of mere species-making as follows:
"Most people of the world, even the most enlightened, are still in the habit of regarding the botanist as a dealer in barbarous Latin names, as a man who gathers flowers, names them, dries them, and wraps them in paper, and all of whose wisdom consists in determining and classifying this hay which he has collected with such great pains."
Although he insisted on correct methods, his ardent nature led him to champion conclusions of his own before they were thoroughly tested. His great influence in the development of scientific botany lay in his earnestness, his application of new methods, and his fearlessness in drawing conclusions, which, although frequently wrong, formed the starting-point of new researches.
Let us now examine the original publications upon which the cell-theory was founded.
Schleiden's Contribution.—Schleiden's paper was particularly directed to the question, How does the cell originate?and was published in Müller'sArchiv, in 1838, under the German title ofUeber Phytogenesis. As stated above, the cell had been recognized for some years, but the question of its origin had not been investigated. Schleiden says: "I may omit all historical introduction, for, so far as I am acquainted, no direct observations exist at present upon the development of the cells of plants."
He then goes on to define his view of the nucleus (cytoblast) and of the development of the cell around it, saying: "As soon as the cytoblasts have attained their full size, a delicate transparent vesicle arises upon their surface. This is the young cell." As to the position of the nucleus in the fully developed cell, he is very explicit: "It is evident," he says, "from the foregoing that the cytoblast can never lie free in the interior of the cell, but is always enclosed in the cell-wall," etc.
Schleiden fastened these errors upon the cell-theory, since Schwann relied upon his observations. On another point of prime importance Schleiden was wrong: he regarded all new cell-formation as the formation of "cells within cells," as distinguished from cell-division, as we now know it to take place.
Schleiden made no attempt to elaborate his views into a comprehensive cell-theory, and therefore his connection as a co-founder of this great generalization is chiefly in paving the way and giving the suggestion to Schwann, which enabled the latter to establish the theory. Schleiden's paper occupies some thirty-two pages, and is illustrated by two plates. He was thirty-four years old when this paper was published, and directly afterward was called to the post of adjunct professor of botany in the University of Jena, a position which with promotion to the full professorship he occupied for twenty-three years.
Schwann's Treatise.—In 1838, Schwann also announced his cell-theory in a concise form in a German scientific periodical, and, later, to the Paris Academy of Sciences; but it was not till 1839 that the fully illustrated account was published. This treatise with the cumbersome title, "Microscopical Researches into the Accordance in the Structure and Growth of Animals and Plants" (Mikroscopische Untersuchungen über die Uebereinstimmung in der Structur und dem Wachsthum der Thiere und Pflanzen) takes rank as one of the great classics in biology. It fills 215 octavo pages, and is illustrated with four plates.
"The purpose of his researches was to prove the identity of structure, as shown by their development, between animals and plants." This is done by direct comparisons of the elementary parts in the two kingdoms of organic nature.
His writing in the "Microscopical Researches" is clear and philosophical, and is divided into three sections, in the first two of which he confines himself strictly to descriptions of observations, and in the third part of which he enters upon a philosophical discussion of the significance of the observations. He comes to the conclusion that "the elementary parts of all tissues are formed of cells in an analogous, though very diversified manner, so that it may be asserted that there is one universal principle of development for the elementary parts of organisms, however different, and that this principle is the formation of cells."
It was in this treatise also that he made use of the term cell-theory, as follows: "The development of the proposition that there exists one general principle for the formation of all organic productions, and that this principle is the formation of cells, as well as the conclusions which may be drawn from this proposition, may be comprised under the termcell-theory, using it in its more extended signification, while, in a more limited sense, by the theory of cells we understand whatever may be inferred from this proposition with respect to the powers from which these phenomena result."
One comes from the reading of these two contributions to science with the feeling that it is really Schwann's cell-theory, and that Schleiden helped by lighting the way that his fellow-worker so successfully trod.
Modification of the Cell-Theory.—The form in which the cell-theory was given to the world by Schleiden and Schwann was very imperfect, and, as already pointed out, it contained fundamental errors. The founders of the theory attached too much importance to the cell-wall, and they described the cell as a hollow cavity bounded by walls that were formed around a nucleus. They were wrong as to the mode of the development of the cell, and as to its nature. Nevertheless, the great truth that all parts of animals and plants are built of similar units or structures was well substantiated. This remained a permanent part of the theory, but all ideas regarding the nature of the units were profoundly altered.
In order to perceive the line along which the chief modifications were made we must take account of another scientific advance of about the same period. This was the discovery of protoplasm, an achievement which takes rank with the advances of greatest importance in biology, and has proved to be one of the great events of the nineteenth century.
The Discovery of Protoplasm and its Effect on the Cell-Theory.—In 1835, before the announcement of the cell-theory, living matter had been observed by Dujardin. In lower animal forms he noticed a semifluid, jelly-like substance, which he designated sarcode, and which he described as being endowed with all the qualities of life. The same semifluid substance had previously caught the attention of some observers, but no one had as yet announced it as the actual living part of organisms. Schleiden had seen it and called it gum. Dujardin was far from appreciating the full importance of his discovery, and for a long time his description of sarcode remained separate; but in 1846 Hugo vonMohl, a botanist, observed a similar jelly-like substance in plants, which he called plantschleim, and to which he attached the name protoplasma.
The scientific world was now in the position of recognizing living substance, which had been announced as sarcode in lower animals, and as protoplasm in plants; but there was as yet no clear indication that these two substances were practically identical. Gradually there came stealing into the minds of observers the suspicion that the sarcode of the zoölogists and the protoplasm of the botanists were one and the same thing. This proposition was definitely maintained by Cohn in 1850, though with him it was mainly theoretical, since his observations were not sufficiently extensive and accurate to support such a conclusion.
Eleven years later, however, as the result of extended researches, Max Schultze promulgated, in 1861, the protoplasm doctrine, to the effect that the units of organization consist of little masses of protoplasm surrounding a nucleus, and that this protoplasm, or living substance, is practically identical in both plants and animals.
The effect of this conclusion upon the cell-theory was revolutionary. During the time protoplasm was being observed the cell had likewise come under close scrutiny, and naturalists had now an extensive collection of facts upon which to found a theory. It has been shown that many animal cells have no cell-wall, and the final conclusion was inevitable that the essential part of a cell is the semifluid living substance that resides within the cavity when a cell-wall is present. Moreover, when the cell-wall is absent, the protoplasm is the "cell." The position of the nucleus was also determined to be within the living substance, and not, as Schleiden had maintained, within the cell-wall. The definition of Max Schultze, that a cell is a globule of protoplasm surrounding a nucleus, marks a new era in the cell-theory, in which the original generalization became consolidated with the protoplasm doctrine.
Further Modifications of the Cell-Theory.—The reformed cell-theory was, however, destined to undergo further modification, and to become greatly extended in its application. At first the cell was regarded merely as an element of structure; then, as a supplement to this restricted view, came the recognition that it is also a unit of physiology,viz., that all physiological activities take place within the cell. Matters did not come to a rest, however, with the recognition of these two fundamental aspects of the cell. The importance of the cell in development also took firmer hold upon the minds of anatomists after it was made clear that both the egg and its fertilizing agents are modified cells of the parent's body. It was necessary to comprehend this fact in order to get a clear idea of the origin of cells within the body of a multicellular organism, and of the relation between the primordial element and the fully developed tissues. Finally, when observers found within the nucleus the bearers of hereditary qualities, they began to realize that a careful study of the behavior of the cell elements during development is necessary for the investigation of hereditary transmissions.
A statement of the cell-theory at the present time, then, must include these four conceptions: the cell as a unit of structure, the cell as a unit of physiological activity, the cell as embracing all hereditary qualities within its substance, and the cell in the historical development of the organism.
Some of these relations may now be more fully illustrated.
Origin of Tissues.—The egg in which all organisms above the very lowest begin, is a single cell having, under the microscope, the appearance shown in Fig. 76. After fertilization, this divides repeatedly, and many cohering cells result. The cells are at first similar, but as they increase in number, and as development proceeds, they grow different, and certaingroups are set apart to perform particular duties. The division of physiological labor which arises at this time marks the beginning of separate tissues. It has been demonstrated over and over that all tissues are composed of cells and cell-products, though in some instances they are much modified. The living cells can be seen even in bone and cartilage, in which they are separated by a lifeless matrix, the latter being the product of cellular activity.
Fig. 76.—The Egg and Early Stages in its Development. (After Gegenbaur.)
Fig. 77 shows a stage in the development of one of the mollusks just as the differentiation of cells has commenced.
The Nucleus.—To the earlier observers the protoplasm appeared to be a structureless, jelly-like mass containing granules and vacuoles; but closer acquaintance with it has shown that it is in reality very complex in structure as well as in chemical composition. It is by no means homogeneous; adjacent parts are different in properties and aptitudes. The nucleus, which is more readily seen than other cell elements,was shown to be of great importance in cell-life—to be a structure which takes the lead in cell division, and in general dominates the rest of the protoplasm.
Chromosomes.—After dyes came into use for staining the protoplasm (1868), it became evident that certain parts of it stain deeply, while other parts stain faintly or not at all. This led to the recognition of protoplasm as made up of a densely staining portion calledchromatin, and a faintly staining portion designatedachromatin. This means of making different parts of protoplasm visible under the microscope led to important results, as when, in 1883, it was discovered that the nucleus contains a definite number of small (usually rod-shaped) bodies, which become evident during nuclear division, and play a wonderful part in that process. These bodies take the stain more deeply than other components of the nucleus, and are designatedchromosomes.
Fig. 77.—An Early Stage in the Development of the Egg of a Rock-Limpet. (After Conklin.)
Attention having been directed to these little bodies, continued observations showed that, although they vary innumber—commonly from two to twenty-four—in different parts of animals and plants, they are, nevertheless, of the same number in all the cells of any particular plant or animal. As a conclusion to this kind of observation, it needs to be said that the chromosomes are regarded as the actual bearers of hereditary qualities. The chromosomes do notshow in resting-stages of the nucleus; their substance is present, but is not aggregated into the form of chromosomes.
Fig. 78.—Highly Magnified Tissue Cells from the Skin of a Salamander in an Active State of Growth. Dividing cells with chromosomes are shown ata,b, andc,. (After Wilson.)
Fig. 78 shows tissue cells, some of which are in the dividing and others in the resting-stage. The nuclei in process of division exhibit the rod-like chromosomes, as shown ata,b, andc.
Fig. 79.—Diagram of the Chief Steps in Cell-division. (After Parker as altered from Fleming.)
Centrosome.—The discovery (1876) of a minute spot of deeply staining protoplasm, usually just outside the nuclearmembrane, is another illustration of the complex structure of the cell. Although the centrosome, as this spot is called, has been heralded as a dynamic agent, there is not complete agreement as to its purpose, but its presence makes it necessary to include it in the definition of a cell.
The Cell in Heredity.—The problems of inheritance, in so far as they can be elucidated by structural studies, have come to be recognized as problems of cellular life. But we cannot understand what is implied by this conclusion without referring to the behavior of the chromosomes during cell-division. This is a very complex process, and varies somewhat in different tissues. We can, however, with the help of Fig. 79, describe what takes place in a typical case. The nucleus does not divide directly, but the chromosomes congregate around the equator of a spindle (D) formed from the achromatin; they then undergo division lengthwise, and migrate to the poles (E,F,G), after which a partition wall is formed dividing the cell. This manner of division of the chromosomes secures an equable partition of the protoplasm. In the case of fertilized eggs, one-half of the chromosomes are derived from the sperm and one-half from the egg. Each cell thus contains hereditary substance derived from both maternal and paternal nuclei. This is briefly the basis for regarding inheritance as a phenomenon of cell-life.
Fig. 80.—Diagram of a Cell. (Modified after Wilson.)
A diagram of the cell as now understood (Fig. 80) will be helpful in showing how much the conception of the cell has changed since the time of Schleiden and Schwann.
Definition.—The definition of Verworn, made in 1895, may be combined with this diagram: A cell is "a body consisting essentially of protoplasm in its general form, including the unmodified cytoplasm, and the specialized nucleus and centrosome; while as unessential accompaniments may be enumerated: (1) the cell membrane, (2) starch grains, (3) pigment granules, (4) oil globules, and (5) chlorophyll granules." No definition can include all variations, but the one quoted is excellent in directing attention to the essentials—to protoplasm in its general form, and the modified protoplasmic parts as distinguished from the unessential accompaniments, as cell membrane and cell contents.
The definition of Verworn was reached by a series of steps representing the historical advance of knowledge regarding the cell. Schleiden and Schwann looked upon the cell as a hollow chamber having a cell-wall which had been formed around the nucleus; it was a great step when Schultze defined the cell in terms of living substance as "a globule of protoplasm surrounding a nucleus," and it is a still deeper level of analysis which gives us a discriminating definition like that of Verworn.
When we are brought to realize that, in large part, the questions that engage the mind of the biologist have their basis in the study of cells, we are ready to appreciate the force of the statement that the establishment of the cell-theory was one of the great events of the nineteenth century, and, further, that it stands second to no theory, with the single exception of that of organic evolution, in advancing biological science.
CHAPTER XII
PROTOPLASM, THE PHYSICAL BASIS OF LIFE
Therecognition of the rôle that protoplasm plays in the living world was so far-reaching in its results that we take up for separate consideration the history of its discovery. Although it is not yet fifty years since Max Schultze established the protoplasm doctrine, it has already had the greatest influence upon the progress of biology. To the consideration of protoplasm in the previous chapter should be added an account of the conditions of its discovery, and of the personality and views of the men whose privilege it was to bring the protoplasm idea to its logical conclusion. Before doing so, however, we shall look at the nature of protoplasm itself.
Protoplasm.—This substance, which is the seat of all vital activity, was designated by Huxley "the physical basis of life," a graphic expression which brings before the mind the central fact that life is manifested in a material substratum by which it is conditioned. All that biologists have been able to discover regarding life has been derived from the observation of that material substratum. It is not difficult, with the help of a microscope, to get a view of protoplasmic activity, and that which was so laboriously made known about 1860 is now shown annually to students beginning biology.
Inasmuch as all living organisms contain protoplasm, one has a wide range of choice in selecting the plant or the animal upon which to make observations.
We may, for illustration, take one of the simplest of animal organisms, the amœba, and place it under the high powersof the microscope. This little animal consists almost entirely of a lump of living jelly. Within the living substance of which its body is composed all the vital activities characteristic of higher animals are going on, but they are manifested in simpler form. These manifestations differ only in degree of development, not in kind, from those we see in bodies of higher organisms.
We can watch the movements in this amœba, determine at first hand its inherent qualities, and then draw up a sort of catalogue of its vital properties. We notice an almost continual flux of the viscid substance, by means of which it is able to alter its form and to change its position. This quality is called that of contractility. In its essential nature it is like the protoplasmic movement that takes place in a contracting muscle. We find also that the substance of the amœba responds to stimulations—such as touching it with a bristle, or heating it, or sending through it a light electric shock. This response is quite independent of the contractility, and by physiologists is designated the property of being irritable.
By further observations one may determine that the substance of the amœba is receptive and assimilative, that it is respiratory, taking in oxygen and giving off carbonic dioxide, and that it is also secretory. If the amœba be watched long enough, it may be seen to undergo division, thus producing another individual of its kind. We say, therefore, that it exhibits the power of reproduction. All these properties manifested in close association in the amœba are exhibited in the bodies of higher organisms in a greater degree of perfection, and also in separation, particular organs often being set apart for the performance of one of these particular functions. We should, however, bear in mind that in the simple protoplasm of the amœba is found the germ of all the activities of the higher animals.
It will be convenient now to turn our attention to the microscopic examination of a plant that is sufficiently transparent to enable us to look within its living parts and observe the behavior of protoplasm. The first thing that strikes one is the continual activity of the living substance within the boundaries of a particular cell. This movement sometimes takes the form of rotation around the walls of the cell (Fig. 81A). In other instances the protoplasm marks out for itself new paths, giving a more complicated motion, called circulation (Fig. 81B). These movements are the result of chemical changes taking place within the protoplasm, and they are usually to be observed in any plant or animal organism.
Fig. 81.—(A) Rotation of Protoplasm in the Cells of Nitella. (B) Highly Magnified Cell of a Tradescantia Plant, Showing Circulation of Protoplasm. (After Sedgwick and Wilson.)
Under the most favorable conditions these movements, as seen under the microscope, make a perfect torrent of unceasing activity, and introduce us to one of the wonderful sights of which students of biology have so many. Huxley(with slight verbal alterations) says: "The spectacle afforded by the wonderful energies imprisoned within the compass of the microscopic cell of a plant, which we commonly regard as a merely passive organism, is not easily forgotten by one who has watched its movement hour by hour without pause or sign of weakening. The possible complexity of many other organisms seemingly as simple as the protoplasm of the plant just mentioned dawns upon one, and the comparison of such activity to that of higher animals loses much of its startling character. Currents similar to these have been observed in a great multitude of very different plants, and it is quite uniformly believed that they occur in more or less perfection in all young vegetable cells. If such be the case, the wonderful noonday silence of a tropical forest is due, after all, only to the dullness of our hearing, and could our ears catch the murmur of these tiny maelstroms as they whirl in the innumerable myriads of living cells that constitute each tree, we should be stunned as with the roar of a great city."
The Essential Steps in Recognizing the Likeness of Protoplasm in Plants and Animals
Dujardin.—This substance, of so much interest and importance to biologists, was first clearly described and distinguished from other viscid substance, as albumen, by Félix Dujardin in 1835. Both the substance and the movements therein had been seen and recorded by others: by Rösel von Rosenhof in 1755 in the proteus animalcule; again in 1772 by Corti in chara; by Mayen in 1827 in Vallisnieria; and in 1831 by Robert Brown in Tradescantia. One of these records was for the animal kingdom, and three were for plants. The observations of Dujardin, however, were on a different plane from those of the earlier naturalists, and heis usually credited with being the discoverer of protoplasm. His researches, moreover, were closely connected with the development of the ideas regarding the rôle played in nature by this living substance.
Dujardin was a quiet modest man, whose attainments and service to the progress of biology have usually been under-rated. He was born in 1801 at Tours, and died in 1860 at Rennes. Being descended from a race of watchmakers, he received in his youth a training in that craft which cultivated his natural manual dexterity, and, later, this assisted him in his manipulations of the microscope. He had a fondness for sketching, and produced some miniatures and other works of art that showed great merit. His use of colors was very effective, and in 1818 he went to Paris for the purpose of perfecting himself in painting, and with the intention of becoming an artist. The small financial returns, however, "led him to accept work as an engineer directing the construction of hydraulic work in Sédan." He had already shown a love for natural science, and this led him from engineering into work as a librarian and then as a teacher. He made field observations in geology and botany, and commenced publication in those departments of science.
About 1834 he began to devote his chief efforts to microscopic work, toward which he had a strong inclination, and from that time on he became a zoölogist, with a steadily growing recognition for high-class observation. Besides his technical scientific papers, he wrote in a popular vein to increase his income. Among his writings of this type may be mentioned as occupying high rank his charmingly written "Rambles of a Naturalist" (Promenades d'un Naturaliste, 1838).
By 1840 he had established such a good record as a scientific investigator that he was called to the newly founded University of Rennes as dean of the faculty. He found himself in an atmosphere of jealous criticism, largely on account of his being elevated to the station of dean, and after two years of discomfort he resigned the deanship, but retained his position as a professor in the university. He secured a residence in a retired spot near a church, and lived there simply. In his leisure moments he talked frequently with the priests, and became a devout Catholic.
His contributions to science cover a wide range of subjects. In his microscopic work he discovered the rhizopods in 1834, and the study of their structure gave him the key to that of the other protozoa. In 1835 he visited the Mediterranean, where he studied the oceanic foraminifera, and demonstrated that they should be grouped with the protozoa, and not, as had been maintained up to that time, with the mollusca. It was during the prosecution of these researches that he made the observations upon sarcode that are of particular interest to us.
His natural history of the infusoria (1841) makes a volume of 700 pages, full of original observations and sketches. He also invented a means of illumination for the microscope, and wrote a manual of microscopic observation. Among the ninety-six publications of Dujardin listed by Professor Joubin there are seven general works, twenty relating to the protozoa, twenty-four to geology, three to botany, four to physics, twenty-five to arthropods, eight to worms, etc., etc. But as Joubin says: "The great modesty of Dujardin allowed him to see published by others, without credit to himself, numerous facts and observations which he had established." This failure to assert his claims accounts in part for the inadequate recognition that his work has received.
Fig. 82.—Félix Dujardin, 1801-1860.
No portrait of Dujardin was obtainable prior to 1898. Somewhat earlier Professor Joubin, who succeeded other occupants of the chair which Dujardin held in the University of Rennes, found in the possession of his descendants aportrait, which he was permitted to copy. The earliest reproduction of this picture to reach this country came to the writer through the courtesy of Professor Joubin, and a copy of it is represented in Fig. 82. His picture bespeaks his personality. The quiet refinement and sincerity of his face areevident. Professor Joubin published, in 1901 (Archives de Parasitologie), a biographical sketch of Dujardin, with several illustrations, including this portrait and another one which is very interesting, showing him in academic costume. Thanks to the spread of information of the kind contained in that article, Dujardin is coming into wider recognition, and will occupy the historical position to which his researches entitle him.
It was while studying the protozoa that he began to take particular notice of the substance of which their bodies are composed; and in 1835 he described it as a living jelly endowed with all the qualities of life. He had seen the same jelly-like substance exuding from the injured parts of worms, and recognized it as the same material that makes the body of protozoa. He observed it very carefully in the ciliated infusoria—in Paramœcium, in Vorticella, and other forms, but he was not satisfied with mere microscopic observation of its structure. He tested its solubility, he subjected it to the action of alcohol, nitric acid, potash, and other chemical substances, and thereby distinguished it from albumen, mucus, gelatin, etc.
Inasmuch as this substance manifestly was soft, Dujardin proposed for it the name of sarcode, from the Greek, meaningsoft. Thus we see that the substance protoplasm was for the first time brought very definitely to the attention of naturalists through the study of animal forms. For some time it occupied a position of isolation, but ultimately became recognized as being identical with a similar substance that occurs in plants. At the time of Dujardin's discovery, sarcode was supposed to be peculiar to lower animals; it was not known that the same substance made the living part of all animals, and it was owing mainly to this circumstance that the full recognition of its importance in nature was delayed.
The fact remains that the first careful studies upon sarcodewere due to Dujardin, and, therefore, we must include him among the founders of modern biology.
Fig. 83.-Purkinje, 1787-1869.
Purkinje.—The observations of the Bohemian investigator Purkinje (1787-1869) form a link in the chain of events leading up to the recognition of protoplasm. Although Purkinje is especially remembered for other scientific contributions, he was the first to make use of the name protoplasm for living matter, by applying it to the formative substance within the eggs of animals and within the cells of the embryo. His portrait is not frequently seen, and, therefore, is included here (Fig. 83), to give a more complete series of pictures of the men who were directly connected with the development of the protoplasm idea. Purkinje was successively a professor in the universities of Breslau and Prague. His anatomical laboratory at Breslau is notable as being one of the earliest (1825) open to students. He went to Prague in 1850 as professor of physiology.
Fig. 84.—Carl Nägeli, 1817-1891.
Von Mohl.—In 1846, eleven years after the discovery of Dujardin, the eminent botanist Hugo von Mohl (1805-1872) designated a particular part of the living contents of the vegetable cell by the term protoplasma. The viscid, jelly-like substance in plants had in the mean time come to be known under the expressive term of plant "schleim." He distinguished the firmer mucilaginous and granular constituent, found just under the cell membrane, from the watery cell-sap that occupies the interior of the cell. It was to the former part that he gave the name protoplasma. Previous to this,the botanist Nägeli had studied this living substance, and perceived that it was nitrogenous matter. This was a distinct step in advance of the vague and indefinite idea of Schleiden, who had in reality noticed protoplasm in 1838, but thought of it merely as gum. The highly accomplished investigator Nägeli (Fig. 84) made a great place for himself in botanical investigation, and his name is connected with several fundamental ideas of biology. To Von Mohl, however, belongs the credit of having brought the word protoplasm into general use. He stands in the direct line of development, while Purkinje, who first employed the wordprotoplasm, stands somewhat aside, but his name, nevertheless, should be connected with the establishment of the protoplasm doctrine.
Fig. 85.—Hugo von Mohl, 1805-1872.
Von Mohl (Fig. 85) was an important man in botany. Early in life he showed a great love for natural science, and as in his day medical instruction afforded the best opportunities for a man with scientific tastes, he entered upon that course of study in Tübingen at the age of eighteen. He took his degree of doctor of medicine in 1823, and spent several years in Munich. He became professor of physiology in Bern in 1832, and three years later was transferred to Tübingen as professor of botany. Here he remained to the end of his life, refusing invitations to institutions elsewhere. He never married, and, without the cares and joys of a family, led a solitary and uneventful life, devoted to botanical investigation.
Cohn.—After Von Mohl's studies on "plant schleim" there was a general movement toward the conclusion that the sarcode of the zoölogists and the protoplasm of the botanists were one and the same substance. This notion was in the minds of more than one worker, but it is perhaps to Ferdinand Cohn (1828-1898) that the credit should be given for bringing the question to a head. After a study of the remarkable movements of the active spores of one of the simplest plants (protococcus), he said that vegetable protoplasm and animal sarcode, "if not identical, must be, at any rate, in the highest degree analogous substances" (Geddes).
Cohn (Fig. 86) was for nearly forty years professor of botany in the University of Breslau, and during his long life as an investigator greatly advanced the knowledge of bacteria. His statement referred to above was made when he was twenty-two years of age, and ran too far ahead of the evidence then accumulated; it merely anticipated the coming period of the acceptance of the conclusion in its full significance.
Fig. 86.—Ferdinand Cohn, 1828-1898.
De Bary.—We find, then, in the middle years of the nineteenth century the idea launched that sarcode and protoplasm are identical, but it was not yet definitely established that the sarcode of lower animals is the same as the living substance of the higher ones, and there was, therefore, lacking an essential factor to the conclusion that there is only one general form of living matter in all organisms. It took another ten years of investigation to reach this end.
The most important contributions from the botanical side during this period were the splendid researches of De Bary (Fig. 87) on the myxomycetes, published in 1859. Here the resemblance between sarcode and protoplasm was brought outwith great clearness. The myxomycetes are, in one condition, masses of vegetable protoplasm, the movements and other characteristics of which were shown to resemble strongly those of the protozoa. De Bary's great fame as a botanist has made his name widely known.
Fig. 87.—Heinrich A. de Bary, 1831-1888.
In 1858 Virchow also, by his extensive studies in the pathology of living cells, added one more link to the chain that was soon to be recognized as encircling the new domain of modern biology.
Fig. 88.—Max Schultze, 1825-1874.
Schultze.—As the culmination of a long period of work, Max Schultze, in 1861, placed the conception of the identitybetween animal sarcode and vegetable protoplasm upon an unassailable basis, and therefore he has received the title of "the father of modern biology." He showed that sarcode, which was supposed to be confined to the lower invertebrates, is also present in the tissues of higher animals, and there exhibits the same properties. The qualities of contractility and irritability were especially indicated. It was on physiological likeness, rather than on structural grounds, that he formed his sweeping conclusions. He showed also that sarcode agreed in physiological properties with protoplasm in plants, and that the two living substances were practically identical. His paper of 1861 considers the living substance in muscles (Ueber Muskelkörperchen und das was man eine Zelle zu nennen habe), but in this he had been partly anticipated by Ecker who, in 1849, compared the "formed contractile substance" of muscles with the "unformed contractile substance" of the lower types of animal life (Geddes).
The clear-cut, intellectual face of Schultze (Fig. 88) is that of an admirable man with a combination of the artistic and the scientific temperaments. He was greatly interested in music from his youth up, and by the side of his microscope was his well-beloved violin. He was some time professor in the University of Halle, and in 1859 went to Bonn as professor of anatomy and director of the Anatomical Institute. His service to histology has already been spoken of (Chapter VIII).
This astute observer will have an enduring fame in biological science, not only for the part he played in the development of the protoplasm idea, but also on account of other extensive labors. In 1866 he founded the leading periodical in microscopic anatomy, theArchiv für Mikroscopische Anatomie. This periodical was continued after the untimely death of Schultze in 1874, and to-day is one of the leading biological periodicals.
It is easy, looking backward, to observe that the period between 1840 and 1860 was a very important one for modern biology. Many new ideas were coming into existence, but through this period we can trace distinctly, step by step, the gradual approach to the idea that protoplasm, the livingsubstance of organism, is practically the same in plants and in animals. Let us picture to ourselves the consequences of the acceptance of this idea. Now for the first time physiologists began to have their attention directed to the actually living substance; now for the first time they saw clearly that all future progress was to be made by studying this living substance—the seat of vital activity. This was the beginning of modern biology.
Protoplasm is the particular object of study for the biologist. To observe its properties, to determine how it behaves under different conditions, how it responds to stimuli and natural agencies, to discover the relation of the internal changes to the outside agencies: these, which constitute the fundamental ideas of biology, were for the first time brought directly to the attention of the naturalist, about the year 1860—that epoch-making time when appeared Darwin'sOrigin of Speciesand Spencer'sFirst Principles.
CHAPTER XIII
THE WORK OF PASTEUR, KOCH, AND OTHERS
Theknowledge of bacteria, those minutest forms of life, has exerted a profound influence upon the development of general biology. There are many questions relating to bacteria that are strictly medical, but other phases of their life and activities are broadly biological, and some of those broader aspects will next be brought under consideration.
The bacteria were first described by Leeuwenhoek in 1687, twelve years after his discovery of the microscopic animalcula now called protozoa. They are so infinitesimal in size that under his microscope they appeared as mere specks, and, naturally, observation of these minute organisms was suspended until nearly the middle of the nineteenth century, after the improvement of microscope lenses. It is characteristic of the little knowledge of bacteria in Linnæus's period that he grouped them into an order, with other microscopic forms, under the namechaos.
At first sight, the bacteria appear too minute to figure largely in human affairs, but a great department of natural science—bacteriology—has been opened by the study of their activities, and it must be admitted that the development of the science of bacteriology has been of great practical importance. The knowledge derived from experimental studies of the bacteria has been the chief source of light in an obscure domain which profoundly affects the well-being of mankind. To the advance of such knowledge we owe the germ-theory of disease and the ability of medical men to cope with contagious diseases. The three greatest names connected with the rise of bacteriology are those of Pasteur, Koch, and Lister, the results of whose labors will be considered later.
Among the general topics which have been clustered around the study of bacteria we take up, first, the question of the spontaneous origin of life.
The Spontaneous Origin of Life
It will be readily understood that the question of the spontaneous generation of life is a fundamental one for the biologist. Does life always arise from previously existing life, or under certain conditions is it developed spontaneously? Is there, in the inorganic world, a happy concourse of atoms that become chained together through the action of the sun's rays and other natural forces, so that a molecule of living matter is constructed in nature's laboratory without contact or close association with living substance? This is a question ofbiogenesis—life from previous life—or ofabiogenesis—life without preëxisting life or from inorganic matter alone.
It is a question with a long history. Its earliest phases do not involve any consideration of microscopic forms, since they were unknown, but its middle and its modern aspect are concerned especially with bacteria and other microscopic organisms. The historical development of the problem may be conveniently considered under three divisions: I. The period from Aristotle, 325B.C., to the experiments of Redi, in 1668; II. From the experiments of Redi to those of Schulze and Schwann in 1836 and 1837; III. The modern phase, extending from Pouchet's observations in 1859 to the present.
I. From Aristotle to Redi.—During the first period, the notion of spontaneous generation was universally accepted, and the whole question of spontaneous origin of life was in a crude and grotesque condition. It was thought that frogsand toads and other animals arose from the mud of ponds and streams through the vivifying action of the sun's rays. Rats were supposed to come from the river Nile, the dew was supposed to give origin to insects, etc.
The scientific writers of this period had little openness of mind, and they indulged in scornful and sarcastic comments at the expense of those who doubted the occurrence of spontaneous generation. In the seventeenth century Alexander Ross, commenting on Sir Thomas Brown's doubt as to whether mice may be bred by putrefaction, flays his antagonist in the following words: "So may we doubt whether in cheese and timber worms are generated, or if beetles and wasps in cow-dung, or if butterflies, locusts, shell-fish, snails, eels, and such life be procreated of putrefied matter, which is to receive the form of that creature to which it is by formative power disposed. To question this is to question reason, sense, and experience. If he doubts this, let him go to Egypt, and there he will find the fields swarming with mice begot of the mud of Nylus, to the great calamity of the inhabitants."
II. From Redi to Schwann.—The second period embraces the experimental tests of Redi (1668), Spallanzani (1775), and Schwann (1837)—notable achievements that resulted in a verdict for the adherents to the doctrine of biogenesis. Here the question might have rested had it not been opened upon theoretical ground by Pouchet in 1859.
The First Experiments.—The belief in spontaneous generation, which was so firmly implanted in the minds of naturalists, was subjected to an experimental test in 1668 by the Italian Redi. It is a curious circumstance, but one that throws great light upon the condition of intellectual development of the period, that no one previous to Redi had attempted to test the truth or falsity of the theory of spontaneous generation. To approach this question from the experimental side was to do a great service to science.
The experiments of Redi were simple and homely. He exposed meat in jars, some of which were left uncovered, some covered with parchment, and others with fine wire gauze. The meat in all these vessels became spoiled, and flies, being attracted by the smell of decaying meat, laid eggs in that which was exposed, and there came from it a large crop of maggots. The meat which was covered by parchment also decayed in a similar manner, without the appearance of maggots within it; and in those vessels covered by wire netting the flies laid their eggs upon the wire netting. There they hatched, and the maggots, instead of appearing in the meat, appeared on the surface of the wire gauze. From this Redi concluded that maggots arise in decaying meat from the hatching of the eggs of insects, but inasmuch as these animals had been supposed to arise spontaneously within the decaying meat, the experiment took the ground from under that hypothesis.
He made other observations on the generation of insects, but with acute scientific analysis never allowed his conclusions to run ahead of his observations. He suggested, however, the probability that all cases of the supposed production of life from dead matter were due to the introduction of living germs from without. The good work begun by Redi was confirmed and extended by Swammerdam (1637-1681) and Vallisnieri (1661-1730), until the notion of the spontaneous origin of any forms of life visible to the unaided eye was banished from the minds of scientific men.
Fig. 89.—Francesco Redi, 1626-1697.
Redi (Fig. 89) was an Italian physician living in Arentino, distinguished alike for his attainments in literature and for his achievements in natural science. He was medical adviser to two of the grand dukes of Tuscany, and a member of the Academy of Crusca. Poetry as well as other literary compositions shared his time with scientific occupations. Hiscollected works, literary, scientific, and medical, were published in nine octavo volumes in Milan, 1809-1811. This collection includes his life and letters, and embraces one volume of sonnets. The book that has been referred to as containing his experiments was entitledEsperienze Intorno Alla Generazione Degl'Insetti, and first saw the light in quarto form in Florence in 1668. It went through five editions in twenty years. Some of the volumes were translated into Latin, and were published in miniature, making books not more than four inches high. Huxley says: "The extreme simplicity of his experiments, and the clearness of his arguments, gained for his views and for their consequences almost universal acceptance."
New Form of the Question.—The question of the spontaneous generation of life was soon to take on a new aspect. Seven years after the experiments of Redi, Leeuwenhoek made known a new world of microscopic organisms—the infusoria—and, as we have seen, he discovered, in 1687, those still minuter forms, the bacteria. Strictly speaking, the bacteria, on account of their extreme minuteness, were lost sight of, but spontaneous generation was evoked to account for the birth of all microscopic organisms, and the question circled mainly around the infusorial animalcula. While the belief in the spontaneous generation of life among forms visible to the unaided eye had been surrendered, nevertheless doubts were entertained as to the origin of microscopic organisms, and it was now asserted that here were found the beginnings of life—the place where inorganic material was changed through natural agencies into organized beings microscopic in size.
More than seventy years elapsed before the matter was again subjected to experimental tests. Then Needham, using the method of Redi, began to experiment on the production of microscopic animalcula. In many of his experiments he was associated with Buffon, the great French naturalist, who had a theory of organic molecules that he wished to sustain. Needham (1713-1784), a priest of the Catholic faith, was an Englishman living on the Continent; he was for many years director of the Academy of Maria Theresa at Brussels. He engaged in scientific investigations in connection with his work of teaching. The results of Needham's first experiments were published in 1748. These experimentswere conducted by extracting the juices of meat by boiling; by then enclosing the juices in vials, the latter being carefully corked and sealed with mastic; by subjecting the sealed bottles, finally, to heat, and setting them away to cool. In due course of time, the fluids thus treated became infected with microscopic life, and, inasmuch as Needham believed that he had killed all living germs by repeated heating, he concluded that the living forms had been produced by spontaneous generation.
Spallanzani.—The epoch-making researches of Spallanzani, a fellow-countryman of Redi, were needed to point out the error in Needham's conclusions. Spallanzani (Fig. 90) was one of the most eminent men of his time. He was educated for the church, and, therefore, he is usually known under the title of Abbé Spallanzani. He did not, however, actively engage in his churchly offices, but, following an innate love of natural science and of investigation, devoted himself to experiments and researches and to teaching. He was first a professor at Bologna, and afterward at the University of Pavia. He made many additions to knowledge of the development and the physiology of organisms, and he was the first to make use of glass flasks in the experimental study of the question of the spontaneous generation of life.