DIATOMACEAE. For the knowledge we possess of these beautiful plants, so minute as to be undiscernible by our unaided vision, we are indebted to the assistance of the microscope. It was not till towards the close of the 18th century that the first known forms of this group were discovered by O. F. Muller. And so slow was the process of discovery in this field of scientific research that in the course of half a century, when Agardh published his Systema algarum in 1824, only forty-nine species included under eight genera had been described. Since that time, however, with modern microscopes and microscopic methods, eminent botanists in all parts of the civilized world have studied these minute plants, with the result that the number of known genera and species has been greatly increased. Over Io,000 species of diatoms have been described, and about 1200 species and numerous varieties occur in the fresh waters and on the coasts of Great Britain and Ireland. Rabenhorst, in the index to his Flora Europaea algarum (1864) enumerated about 4000 forms which had up to that time been discovered throughout the continent of Europe.
The diatoms are more commonly known among systematic botanists as the Bacillarieae, particularly on the continent of Europe, and although such an immense number of very diverse forms are included in it, the group as a whole exhibits a remarkable uniformity of structure. The Bacillarieae is one of the large groups of Algae, placed by some in close proximity to the FIG. I.
A and B, Melosira arenaria. C-E, Melosira varians. E, showing formation of auxospore. All X450.
Conjugatae and by others as an order of the Brown Algae (or Phaeophyceae); but their characters are so distinctive and their structure is so uniform as to warrant the separation of the diatoms as a distinct class. The affinities of the group are doubtful.
The diatoms exhibit great variety of form. While some 2. - Synedra Ulna. X200. species are circular and more or less disk-shaped, others are oval in outline. Some are linear, as Synedra Ulna (fig. 2); others more or less cres centic; others again are cuneate, as Podosphenia Lyngbyii (fig. 3); some few have a sigmoid outline, as Pleurosigma balticum (fig. 4); but the prevailing forms are naviculoid, as in the large family Naviculaceae, of which the genus Navicula embraces upwards of 1000 species. They vary also in their modes of growth, - some being free-floating, others attached to foreign bodies by simple or branched gelatinous stalks, which in some species are short and thick, while in others they are long and slender. In some genera the forms are simple, while in others the frustules are connected together in ribbon-like filaments, or form, as in other cases, zigzag chains. In some genera the individuals are naked, while in many others they are enclosed in a more or less definite gelatinous investment. The conditions necessary to their growth are moisture and light. Wherever these circumstances coexist, diatomaceous forms will almost invariably be found. They occur mixed with other organisms on the surface of moist rocks; in streamlets and pools, they form a brownish stratum on the surface of the mud, or cover the stems and leaves of water plants or floating twigs with a furry investment. Marine forms are usually attached to various sea-weeds, and many are found in the stomachs of molluscs, holothurians, ascidians and other denizens of the ocean. The fresh-water forms are specifically distinct from those incidental to salt or brackish water, - fresh-water species. however, are sometimes FIG. 5.
A-C, Tetracyclus lacustris. D and E, Tabellaria fenestrata. F and G, Tabellaria flocculosa. All X 500.
carried some distance into the sea by - the force of the current, and in tidal rivers marine forms are carried up by the force of the tide. Some notion may be formed of the extreme minuteness of these forms from the fact that one the length of which is 2 u?th of an inch may be considered as beyond the medium size. Some few, indeed, are much larger, but by far the greater proportion are of very much smaller dimensions.
Diatoms are unicellular plants distinguished from kindred forms by the fact of having their soft vegetative part covered by a siliceous case. Each individual is known as a frustule, and the cell-wall consists of two similar valves nearly parallel to each other, each valve being furnished with a rim (or connecting-band) projecting from it at a right angle.
One of these valves with its rim is slightly smaller than the viii. 6 a FIG. 3. - Podosphenia Lyngbyii. X400.
? ?:;: ?-?;,; ,??;?. FIG. 4. - Pleurosigma balticum. X200.
other, the smaller fitting into the larger pretty much as a pill-box fits into its cover. This peculiarity of structure affords ample scope for the growth of the protoplasmic cell-contents, for as the latter increase in volume the siliceous valves are pushed out, and their corresponding siliceous rims become broader. The connecting-bands although closely fitting their respective valves are distinct from them, and together the two bands form the girdle.
An individual diatom is usually described from two aspects, one in which the surface of the valve is exposed to view - the valve view, and one in which the girdle side is exposed - the girdle view. The valves are thin and transparent, convex on the outside, and generally ornamented with a variety of sculptured markings. These sculptures often present the aspect of striae across the face of the valve, and the best lenses have shown them to .consist of a series of small cavities within the siliceous wall of the cell. The valves of some of the marine genera exhibit a beautiful areolated structure due to the presence of larger chambers within the siliceous cell-wall. Many diatoms possess thickenings of the cell-wall, visible in the valve view, in the centre of the valve and at each extremity. These thickenings are known as the nodules, and they are generally connected by a long median line, the raphe, which is a cleft in the siliceous valve, extending at least some part of its length.
The protoplasmic contents of this siliceous box-like unicell are very similar to the contents of many other algal cells. There is a living protoplasmic layer or primordial utricle, connected either by two broad bands or by a number of anastomosing threads with a central mass of protoplasm in which the nucleus is embedded. The greater part of the cavity of the cell is occupied by one or several fluid vacuoles. The characteristic brown colour of diatoms is due to the presence of chromatophores embedded in the lining layer of protoplasm. In number and form these chromatophores are variable. They contain chlorophyll, but the green colour is masked by the presence of diatomin, a brown pigment which resembles that which occurs in the Brown Algae or Phaeophyceae. The chromatophores contain a variable number of pyrenoids, colourless proteid bodies of a crystalloidal character.
One of the first phenomena which comes under the notice of the observer is the extraordinary power of motion with which the frustules are endowed. Some species move slowly backwards and forwards in pretty much the same line, but in the case of Bacillaria paradoxa the motion is very rapid, the frustules darting through the water in a zigzag course. To account for this motion various theories have been suggested, none of which appear to be altogether satisfactory. There is little doubt that the movements are connected with the raphe, and in some diatoms there is much evidence to prove that they are due to an exudation of mucilage.
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The most natural system of classification of the Bacillarieae is the one put forward by Schutt (1896), and since generally followed by systematists. He separates them into two primary divisions, the ` Centricae ' and the ` Pennatae.' The former includes all those diatoms which in the valve view possess a radial symmetry around a central point, and which are destitute of a raphe (or a pseudoraphe). The latter includes those which are zygomorphic or otherwise irregular, and in which the valve view is generally boat-shaped or needle-shaped, with the markings arranged in a sagittal manner on each side of a raphe or pseudoraphe.
In the Diatomaceae, as well as in the Desmidieae, the ordinary mode of increase is by simple cell-division. The cell-contents within the enclosure of the siliceous case separate into two distinct masses. As these two daughter-masses become more and more developed, the valves of the mother-cell are pushed more and more widely apart. A new siliceous valve is secreted by each of the two masses on the side opposite to the original valve, the new valves being situated within the girdle of the original frustule. When this process has been completed the girdle of the mother frustule gives way, and two distinct frustules are formed, the siliceous valves in each of these new frustules being one of the valves of the mother-cell, and a newly formed valve similar and more or less parallel to it.
During the life of the plant this process of self-division is continued with an almost incredible rapidity. On this subject the observation of Professor William Smith, writing in 1853, is worthy of special notice: - "I have been unable to ascertain the time occupied in a single act of self-division, but supposing it to be completed in twenty-four hours we should have, as the progeny of a single frustule, the amazing number of r,000,000,000 in a single month, a circumstance which will in some degree explain the sudden, or at least rapid, appearance of these organisms in localities where they were a short time previously either unrecognized or sparingly diffused" (British Diatomaceae, vol. i. p. 25).
Individual diatoms when once produced by cell-division are incapable of any increase in size owing to the rigidity of their siliceous cell-walls, and since the new valves are always formed within the girdle of the old ones, it would follow that every succeeding generation is reduced in size by the thickness of the girdle. In some diatoms, however, this is not strictly true as daughter-cells are sometimes produced of larger size than the parent-cells. Thus, the reduction in size of the individuals is not always proportionate to the number of cell-divisions.
On the diminution in size having reached a limit in any species, the maximum size is regained by the formation of an auxospore. There are five known methods of reproduction by auxospores, but it is unnecessary here to enter into details of these methods. Suffice it to say that a normal auxospore is produced by the conjugation of two parent-cells, its distinguishing feature being a rejuvenescence accompanied by a marked increase in size. These auxospores formed without conjugation are parthenogenetic.
The Diatomaceae are usually gathered in small bottles, and special care should be taken to collect them as free as possible from extraneous matter. A small portion having been examined under the microscope, should the gathering be thought worthy of preservation, some of the material is boiled in acid for the purpose of cleaning it. The acids usually employed are hydrochloric, nitric or sulphuric, according as circumstances require. When the operator considers that by this process all foreign matter has been eliminated, the residuum is put into a precipitating jar of a conical shape, broader at the bottom than at the top, and covered to the brim with filtered or distilled water. When the diatoms have settled in the bottom of the jar, the supernatant fluid is carefully removed by a syringe or some similar instrument, so that the sediment be not disturbed. The jar is again filled with water, and the process repeated till the acid has been completely removed. It is desirable afterwards to boil the sediment for a short time with supercarbonate of soda, the alkali being removed in the same manner as the acid. A small portion may then be placed with a pipette upon a slip of glass, and, when the moisture has been thoroughly evaporated, the film that remains should be covered with dilute Canada balsam, and, a thin glass cover having been gently laid over the balsam, the preparation should be laid aside for a short time to harden, and then is ready for observation.
Diatoms are most abundant in cold latitudes, having a general preference for cold water. In the pelagic waters of lakes and of the oceans they are often very abundant, and in the cold waters of the Arctic and Antarctic C FIG. 6. - Formation of Auxospores.
A. Navicula limosa. B. Achnanthes flexella C. Navicula Amphisbaena. D. Navicula viridis. A-C, X 450; D, X;350.
Oceans they exist in prodigious numbers. They thus form a large proportion of both the marine and the fresh-water plankton.
Large numbers of fossil diatoms are known. Not only are these minute plants assisting at the present time in the accumulation of oceanic and lake deposits, but in former ages they have been sufficiently active to give rise to considerable deposits of diatomaceous earths. When the plant has fulfilled its natural course the siliceous covering sinks to the bottom of the water in which it had lived, and there forms part of the sediment. When in the process of ages, as it has often happened, the accumulated sediment has been hardened into solid rock, the siliceous frustules of the diatoms remain unaltered, and, if the rock be disintegrated by natural or artificial means, may be removed from the enveloping matrix and subjected to examination under the microscope. The forms found may from their character help in some degree to illustrate the conditions under which the stratum of rock had been originally deposited. These earths are generally of a white or grey colour. Some of them are hard, but most are soft and friable. Many of them are of economic importance, being used as polishing powders ("Tripoli"), as absorbents for nitroglycerin in the manufacture of dynamite ("Kieselguhr"), as a dentifrice, and more recently they have been used to a large extent in the manufacture of non-conducting and sound-proof materials. Most of these diatomaceous earths are associated with rocks of Tertiary formations, although it is generally regarded that the earliest appearance of diatoms is in the Upper Cretaceous (chalk).
Vast deposits of Diatomaceous earths have been discovered in various parts of the world, - some the deposit of fresh, others of salt water. Of these deposits the most remarkable for extent, as well as for the number and beauty of the species contained in it, is that of Richmond, in Virginia, one of the United States of America. It extends for many miles, and is in some places at least 40 ft. deep. It is a remarkable fact that though the generations of a diatom in the space of a few months far exceed in number the generation of man during the period usually assigned to the existence of the race, the fossil genera and species are in most respects to the most minute details identical with the numerous living representatives of their class.
(E. O'M.; G. S. W.*)
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