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See also:PROTOZOA (Gr. 7rpwros, first, and i'woe, living thing) , the name given by See also:modern zoologists to the animalcules, for the most See also:part microscopic, which were termed by the older naturalists See also:Infusoria, from the manner in which they appear in infusions containing decaying See also:animal and See also:vegetable See also:matter . The name Infusoria is now, however, restricted to one of the four classes which comprise the See also:Protozoa proper . The name Protozoa was coined as far back as 182o as an See also:equivalent for the See also:German word Urthiere, meaning animals of See also:primitive or archaic nature, the forms of animal See also:life which may be supposed to have been the first that appeared upon our globe . The See also:great naturalist C . T. von See also:Siebold was, however, the first to give a scientific See also:definition to the See also:group . Von Siebold pointed out that in the Protozoa the individual was always a single vital unit or See also:cell, in contrast with the higher See also:division of the animal See also:kingdom, the Metazoa, in which the See also:body is generally, though not universally, regarded as composed of many such See also:units . To put the matter briefly and somewhat technically: the Protozoa are unicellular animals, the Metazoa multicellular animals; in the Protozoa the cell is See also:complete in itself, both morphologically and physiologically, and is capable of maintaining a See also:separate and See also:independent existence in suitable surroundings, like any other organism; in the Metazoa the cells are differentiated for the performance of distinct functions and combined together to See also:form the various tissues of which the body is built up, and the individual cells of the Metazoan body are not capable of maintaining a separate existence apart from their See also:fellows . This is the sense in which the See also:term Protozoa is used by zoologists, whereby certain forms of animal life, which were formerly ranked as Protozoa, such as See also:sponges and rotifers, are now definitely excluded from the group and classed as Metazoa . The animal kingdom may be divided, therefore, into two sub-kingdoms, the Protozoa and the Metazoa, the first-named characterized by their essentially unicellular nature . This is a criterion by which it is easy to define the Protozoa from a purely zoological standpoint, but which becomes less satisfactory when we take into See also:consideration the whole range of microscopic unicellular organisms . Besides the true Protozoa, which, ex hypothesi, are organisms of animal nature, there are many other organisms of equally See also:simple organization, including the Bacteria and the unicellular See also:plants . The Bacteria stand sharply apart from the other forms of life, not only, in many cases, by their divergent methods of See also:metabolism, but by morphological characteristics, such as the definite body-form limited by a distinct envelope, the See also:absence of See also:organs for locomotion other than the See also:peculiar flagella, and, above all, by the lack of any differentiation of the body-See also:protoplasm into See also:nucleus and cytoplasm, as in all true cells of either animal or vegetable nature . On the other See also:hand, to separate by hard-and-fast See also:definitions the unicellular plants from the unicellular animals is not only difficult but practically impossible . The essential difference between plant and animal is a physiological one, a difference in the method of See also:nutrition . A typical See also:green plant is able to live independently of other organisms and to build up its substance from simple gases in the See also:air and inorganic salts in the See also:soil or See also:water, provided that certain conditions of See also:light and moisture be See also:present in its environment; this is the so-called holophytic method of nutrition . A typical animal, on the other hand, while practically independent of sunlight, is not able to exist apart from other living organisms, since it is not able to build up its substance from simple chemical constituents like a plant, but must be supplied with ready-made proteids in its See also:food, for which it requires other organisms, either plants or animals; this is the so-called holozoic method of nutrition . Intermediate between these two habits of life is the so-called saprophytic See also:habit, exemplified by the See also:fungi amongst plants; in this method of nutrition the organism cannot build up its substance entirely from inorganic substances, but absorbs the organic substances present in solutions containing organic salts or decaying animal or vegetable matter . If we regard the organisms termed collectively Protozoa from the point of view of their methods of nutrition (considering for the present only See also:free-living, non-parasitic forms), we find in one class, the See also:Flagellata, examples of the three methods mentioned above, the holozoic, holophytic and saprophytic habit of life, not only in See also:species closely allied to each other, but even combined in one and the same species at different periods of its life or in different surroundings . An individual of a given species may contain See also:chlorophyll, with which it decomposes carbonic See also:acid See also:gas in the sunlight, like a plant, while possessing a definite mouth-See also:aperture, by means of which it can ingest solid food, like an animal . Such instances show clearly that in the simplest forms of life the difference between plant and animal is but a difference of habit and of mode of nutrition, to which the organism is not at first irrevocably committed, and which are not at first accompanied by distinctive morphological characteristics . Only when the organism becomes specialized for one or the other mode of life exclusively does it acquire such definite morphological characters that the difference between plant and animal can be used for the purpose of a natural See also:classification, as in the higher forms of life . In the lowest forms it is not possible to See also:base natural subdivisions on their vegetable or animal nature . - For this See also:reason it has been proposed by E . See also:Haeckel to unite all the primitive forms of life in which the body is morphologically equivalent to a single cell into one group, the See also:Protista, irrespective of their animal or vegetable nature . In this method of dealing with the problem the Protista are regarded as a distinct kingdom (Reich), more or less inter-mediate between, but distinct from, the animal and vegetable kingdoms, and representing the ancestral stock from which both animals and plants have sprung . Many authorities have followed Haeckel's See also:lead in the matter, and the See also:science of Protistology or Protistenkunde has already a See also:special See also:journal devoted to the publication of researches upon it . But though it may be more scientific, from a theoretical point of view, to group all these primitive organisms together in the way suggested by Haeckel, in practice it is inconvenient, on See also:account of thevast number of forms of life to be comprised as Protista, their diversity in habit of life and organization, and, above all, the difference in the technical methods required for their study, which becomes too complicated for a single worker . Hence Protistology becomes split up in practice by its own See also:mass into three sciences: the Bacteria are the See also:objects of the science of See also:bacteriology; botanists See also:deal with the unicellular plants; and the zoologists with those Protista which are more distinctly animal in their characters . Hence the Protozoa are to be regarded as a convenient rather than a natural group, and may be characterized generally as follows: Organisms in which the individual is a single cell, that is to say, consists of a single undivided mass of protoplasm which is capable of independent existence in a suitable environment; if many such individuals be combined together to form a See also:colony, as frequently occurs, there is no differentiation of the individuals except for reproductive purposes, and never for See also:tissue-formation as in the Metazoa . The body always contains chromatin or nuclear substance, which may be disposed in various ways, but usually forms one or more concentrated masses termed nuclei, which can be distinguished sharply from the See also:general body-protoplasm or cytoplasm . The protoplasmic body may be naked at the See also:surface, or maybe limited and enclosed by a distinct envelope or cell-membrane, which is not usually of the nature of See also:cellulose, except in holophytic forms . Organs serving for locomotion and for the See also:capture and assimilation of solid food are usually present, but may be wanting altogether when the mode of nutrition is other than holozoic; chlorophyll, on the other hand, is only found as a constituent of the body-substance in the holophytic Flagellata.' To these characters it may be added that See also:reproduction is effected by some form of fission, or division of the body into smaller portions, and that in the vast See also:majority of Protozoa, if not in all, a See also:process of conjugation or syngamy occurs at some See also:period in the life-See also:cycle, the essential feature of the process being See also:fusion of nuclear matter from distinct individuals . The foregoing definition does not distinguish the Protozoa sharply from the primitive forms of plant-life, with which, as stated above, they are connected by many transitions; but the differentiation of the body-substance into nucleus and cytoplasm separates them at once from the Bacteria, in which the chromatin is distributed evenly through the body-protoplasm . Protozoa and Disease.—The study of the Protozoa has acquired great See also:practical importance from the fact that many of them live as parasites of other animals, and as such may be the cause of dangerous diseases and epidemics in the higher forms of animal life and in See also:man (see PARASITIC DISEASES) . Examples of parasitic forms are to be found in all the four classes into which, as will be stated below, the Protozoa are divided, and one class, the See also:Sporozoa, is composed entirely of endoparasitic forms . Hence Protozoology, as it is termed, is rapidly assuming an importance in medical and veterinary science almost equal to that of bacteriology, although the recognition of Protozoa as agents in the See also:production of disease is hardly older than a See also:decade .
The most striking instances of Protozoa well established as pathogenic agents are the malarial parasites, the species of Piro plasma causing haemoglobinuria of See also:cattle and other animals, the See also:trypanosomes causing tsetse-See also:fly disease, surra, sleeping' sickness, and other maladies, the species of Leishmania causing kala azar and See also:oriental sore, and the See also:Amoeba responsible for the so-called amoebic See also:dysentery
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Other diseases referred, but as yet doubtfully, to the agency of Protozoa are syphilis, small-pox, See also:hydrophobia, yellow See also:fever, and even See also:cancer
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It is only possible here to discuss briefly in a general way the relations of these parasites to their hosts
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When two organisms stand habitually in the relation of See also:host and See also:parasite, an See also:equilibrium tends to become established gradually between them, so
1 Many Protozoa contain symbiotic green organisms, so-called zoochlorellae or zooxanthellae, in their body-protoplasm; for instance, See also:Radiolaria, and See also:Ciliata such as See also:Paramecium bursaria, &e
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This See also:condition must be carefully distinguished from chlorophyll occurring as a cell-constituent
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that a condition is brought about in which, after many generations, the host becomes " tolerant " of the parasite, and the parasite is not lethal to the host, though perhaps capable of setting up considerable disturbance in its vital functions
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Many animals are found to contain almost constantly certain See also:internal parasites without being, apparently, in the least affected by them; and it should be See also:borne in mind that in most cases it is not to the See also:interest of the parasite to destroy the host or to over-tax its resources
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But when the parasite is transferred naturally or artificially to a species or See also:race of host which does not ordinarily See also:harbour it, and which therefore has not acquired See also:powers of resisting its attacks, the parasites may be most deadly in their effects
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Thus the See also: The transference may take See also:place naturally, by the bite of a tsetse-fly, or may be effected artificially; in either See also:case T. brucii is extremely lethal to certain animals, such as imported cattle, horses and See also:dogs, or to rats and See also:guinea-pigs . Other animals, however, may be quite " repellent "I to this parasite, that is to say, if it be inoculated into their See also:blood it See also:dies out without producing See also:ill effects, just as T. lewisi does when injected into an animal other than a rat . Thus it is seen that T. brucii, when introduced into the blood of an animal which is specifically or racially distinct from its natural hosts in the region where it is indigenous, is either unable to maintain itself in its new host, or flourishes in it to such an extent as to be the cause of its See also:death . We may assume, therefore, at least as a working See also:hypothesis, that a lethal parasite is one that is new to its host, and that a harmless parasite is one See also:long established . Since all parasites must have been new to their proper hosts at some period, See also:recent or remote, in the See also:history of the species, it would follow that the first commencement of See also:parasitism would be in almost all cases a life and death struggle, as it were, between the two organisms concerned, and it is quite conceivable that the host might succumb in the struggle and so be exterminated . See also:Ray Lankester has suggested that the extinction of many species of animals in the past may have been due, in some cases, to their having been attacked by a species of parasite to which they did not succeed in becoming adapted, and by which they became, in consequence, exterminated entirely . Organization of the Protozoa.—The body-form may be See also:constant or inconstant in the Protozoa, according as the body-substance is or is not limited at the surface by a See also:firm envelope or cuticle . When the surface of the protoplasm is naked, as in the See also:common amoeba and allied organisms, the movements of the animal bring about continual changes of form . The protoplasm flows out at any point into processes termed pseudopodia, which are being continually retracted and formed anew . Such movements are known as amoeboid, and may be seen in the cells of Metazoa as well as in Protozoa . The pseudopodia serve both for locomotion and for the capture of food . If equally See also:developed on all sides of the body, the animal as a whole remains stationary, but if formed more on one See also:side than the other, the mass of the body shifts its position in that direction, but the See also:movement of See also:translation is generally slow . If the animal remains perfectly quiescent and inactive, the See also:laws of surface-tension acting upon the semi-fluid protoplasmic body cause it to assume a simple spherical I The use of the terms " tolerant " and " repellent " is taken from the excellent See also:article on " Sleeping Sickness," by E . Ray Lankester, in the Quarterly See also:Review (See also:July 1904), No . 399. pp . 113-138 . In the majority of Protozoa, however, the protoplasm is limited at the surface by a firm membrane or cuticle, and in consequence the body has a definite form, which varies greatly in different species, according to the habit of life . As a general rule those forms that are fixed and sedentary in habit tend towards a radially symmetrical structure; those that are free-See also:swimming approach to an ovoid form, with the longest See also:axis of the body placed in the direction of movement; and those that creep upon a firm substratum have the See also:lower side of the body flattened, so that dorsal and ventral surfaces can be distinguished; it is very rare, however, to find a bilaterally symmetrical type of body-structure amongst these organisms . In some cases the cuticle may be too thin to check completely the changes of form due to the movements of the underlying protoplasm; instances of this are seen amongst the so-called " metabolic " Flagellata, in which the body exhibits continually changes of form, termed by Lankester " euglenoid " movements, due to the activity of the superficial contractile layer of the body manifesting itself in See also:ring-like contractions passing down the body in a manner similar to the peristaltic movements of the See also:intestine . The body-substance of the Protozoa is protoplasm, or, as it was originally termed by Dujardin, sarcode, which is finely alveolar in structure, the See also:diameter of the alveoli varying generally between z and r µ . At the surface of the body the alveoli may take on a definite See also:honeycomb-like arrangement, forming a special " alveolar layer " which in See also:optical See also:section appears radially striated . Besides the See also:minute protoplasmic alveoli, the protoplasm often shows a coarse vacuolation throughout the whole or a part of its substance, giving the body a frothy structure . When such vacuoles are present they must be carefully distinguished from the contractile vacuoles and food-vacuoles described below; from the former they differ by their non-contractile nature, and from the latter by not containing food-substances . In many Protozoa and especially in those forms in which there is no cuticle, the body may be supported by a See also:skeleton . The material of the skeleton differs greatly in different cases, and may be wholly of an organic nature, or may be impregnated with, or almost entirely composed of, inorganic See also:mineral salts, in which case the skeletal substance is usually either See also:silica or carbonate of See also:lime . From the morphological point of view the skeletons of Protozoa may be divided into two See also:principal classes, according as they are formed internal to, or See also:external to, the body in each case . Instances of internal skeletons are best seen in the spherical floating forms comprised in the orders Radiolaria and See also:Heliozoa; such skeletons usually take the form of spicules, radiating from the centre to the circumference, and often further strengthened by the for. mation of tangential bars, producing by their See also:union a lattice-See also:work, which in species of relatively large See also:size may be formed periodically at the surface as the animal grows so that the entire skeleton takes the form of concentric hollow See also:spheres held together by radiating beams . The architectural types of these skeletons show, however, an almost See also:infinite diversity, and cannot be summarized briefly . External skeletons have usually the form of a See also:shell or See also:house, into which the body can be retracted for See also:protection, and from which the protoplasm can issue forth during the animal's phases of activity . Shells of this See also:kind, which must be carefully distinguished from cuticles or other membranes that invest the body closely, are well seen in the See also:order See also:Foraminifera; in the simplest cases they are monaxon in See also:architecture, that is to say, with one principal axis See also:round which the shell is radially symmetrical, and at one See also:pole is a large aperture through which the protoplasm can creep out . In addition to the principal aperture, the shell may or may not be pierced all over by numerous See also:fine pores, through which also the protoplasm can pass out . For further details concerning these shells and their very numerous varieties of structure the reader is referred to the article FORAMINIFERA . The protoplasmic body of the Protozoa is frequently differ- of the nuclear apparatus set apart as a distinct kinetic nucleus, entiated into two zones or regions: a more external, termed the with the See also:function, apparently, of governing the activities of ectoplasm or ectosarc, and a more internal, termed the endo- the flagellum . plasm or endosarc . The ectosarc is distinguished by being See also:Cilia are minute, See also:hair-like extensions of the ectoplasm, which more clear and hyaline in See also:appearance, and more tough and viscid See also:pierce the cuticle and form typically a furry covering to the body. in consistence; the endoplasm, on the other hand, is more I Though perhaps primitively derived from flagella, cilia, in their granular and opaque, and of a more fluid nature . The ecto- usual form, are distinguished from flagella by being of smaller plasm is the protective layer of the body, and is also the portion size, by being present, as a rule, in much greater See also:numbers, and most concerned in movement, in See also:excretion, and perhaps also above all by the See also:character of their movements . In the place in sensation and in functions similar to those performed by the of the complicated lashing movements of the flagella, each cilium See also:nervous systems of higher animals . The endoplasm, on the performs a simple stroke in one direction, becoming first bowed other hand, is the See also:chief seat of See also:digestive and reproductive on one side, by an See also:act of contraction, and then straightened functions. out again when relaxed . The movements of the cilia are co- As the protective layer of the body, the ectoplasm forms ordinated and they act in See also:concert, though not absolutely in the envelopes or membranes which invest the surface of the body, unison, each one contracting just before or after its See also:neighbour, and which are differentiations of the outermost layer of the so that waves of movement pass over a ciliated surface in a ectoplasm . Thus in most Flagellata the ectoplasm is represented given direction, similar to what may be seen in a cornfield when only by the more or less firm See also:outer covering or periplast . Even the See also:wind is blowing over it . Primitively coating the whole when such envelopes are absent, however, the ectoplasm can surface of the body evenly, the'cilia may become modified and still be seen to exert a protective function; as, for instance, in specialized in various ways, which cannot be described in detail those Myxosporidia which are parasitic in the See also:gall-bladders or here (see INFUSORIA) . urinary bladders of their hosts, and which can resist the See also:action Besides the organs of locomotion already mentioned, there of the juices in which they live so long as the ectoplasm is intact, may be present so-called undulating membranes, in the form but succumb to the action of the See also:medium if the ectoplasm be of thin sheets of ectoplasm which are capable of performing injured . In many Infusoria the ectoplasm contains special sinuous, undulating movements by their inherent contractility. organs of offence termed trichocysts, each a minute ovoid body In some cases distinct contractile threads or See also:myonemes have from which, on stimulation, a See also:thread is shot out, in a manner been described in these membranes . Undulating membranes similar to the nematocysts of Coelenterata . Similar organs I appear to be formed either by the fusion together of a See also:row of are seen also in the spores of Myxosporidia, as the so-called polar cilia, side by side,. or by the See also:attachment of a flagellum to the capsules; but in this case the organs are not specially ectoplasmic, body by means of an ectoplasmic See also:web, in which case the flageland appear to serve for See also:adhesion and attachment, rather than lum forms the free edge of the membrane, as in the genus for offence . Trypanosoma . The connexion of the ectoplasm with movement is seen in the Returning to the ectoplasm, the excretory function exerted simplest forms, such as Amoeba, by the fact that all pseudopodia by this layer is seen by the formation in it of the peculiar See also:con-arise from it in the first instance . In forms with a definite tractile vacuoles found in most free-living Protozoa . A con- cuticle, on the other hand, the ectoplasm usually contains contractile See also:fibres or myonemes, forming, as it were, the See also:muscular See also:system of the organism . The dependence of the motility of the animal upon the development of the ectoplasm is well seen in See also:Gregarines, in which other organs of locomotion are absent; in forms endowed with active powers of locomotion a distinct limit of its growth it discharges its contents to the exterior by ectoplasmic layer is present below the cuticle; in those Gregarines a sudden and rapid contraction . There is, apparently, in most incapable of active movement, on the other hand, the ectoplasm if not in all cases, a definite See also:pore through which the contractile vacuole empties itself to the exterior . On account of the relatively large size which the contractile vacuole attains it bulges inwards beyond the limits of the ectoplasm and comes to See also:lie chiefly in the endoplasm, to which it is sometimes, but erroneously, ascribed . In the most highly differentiated Protozoa, for instance, the Ciliata, the ectoplasm contains an apparatus of excretory channels, situated in its deeper layers, and forming as it were a drainage-system, from which the contractile vacuoles are fed . The fluid discharged by the contractile vacuoles appears to be chiefly water which has been absorbed at the surface of the protoplasmic body, and which has filtered through the protoplasm, taking up the soluble See also:waste nitrogenous products of the metabolism and the gaseous products of respiration; hence the contractile vacuoles may be compared in a general way to the urinary and See also:respiratory organs of the Metazoa . One of the first consequences of the parasitic habit of life is the disappearance of the contractile vacuoles, which are hardly ever found in truly parasitic Protozoa, that is to say, in forms which live in the interior of other animals and nourish them-selves at their expense . They are also very frequently absent in marine forms . Mechanisms of a nervous nature are very seldom found in Protozoa, but in some Ciliata special tactile bristles are found, and it is possible that flagella, and perhaps even pseudopodia, may be sometimes tactile rather than locomotor in function . Pigment-spots, apparently sensitive to light, may also occur in some Flagellata . The endoplasm, as already stated, is the chief seat of nutritive and reproductive processes . In many Flagellata the ectoplasm tractile vacuole is a spherical drop of watery fluid which makes its appearance periodically at some particular spot near the surface of the animal's body, or, if more than one such vacuole is present, at several definite and constant places . Each vacuole grows to a certain size, and when it has reached the is absent or scarcely recognizable . From the ectoplasm arise the special organs of locomotion, which, when present, take the form of pseudopodia, flagella or cilia . Pseudopodia, as already explained, are temporary protoplasmic organs which can be extruded or retracted at any point; they fall naturally into two principal types, between which, however, transitions are to be found: first, slender, filamentous or filose pseudopodia, composed of ectoplasm alone, which may remain separate from one another, or may anastomose to form networks, and are then termed reticulose; secondly, thick, See also:blunt, so-called lobose pseudopodia, which are composed of ectoplasm with a core of endoplasm, and never form networks . In forms showing active locomotor powers the pseudopodia are usually more lobose in type; filose pseudopodia, on the other hand, are more adapted for the function of capturing food . Flagella are long, slender, vibratile filaments, generally few in number when present, and usually placed at the pole of the body which is anterior in progression . Each flagellum performs peculiar lashing movements which cause the body, if free, to be dragged along after the flagellum in jerks or leaps; if, however, the body be fixed, the action of the flagellum or flagella causes a current towards it, by which means the animal obtains its food-See also:supply . A flagellum which is anterior in movement has been distinguished by Lankester by the convenient term tractellum; sometimes, however, the flagellum is posterior in movement and acts as a propeller, like the tail of a See also:fish; for this type Lankester has proposed the term pulsellum . The flagellum appears to arise in all cases from a distinct basal granule, and in some cases, as in the genus Trypanosoma, there is a portion is represented only by the thin envelope or periplast, so that the whole body is practically endoplasm . When the two layers are well differentiated the endoplasm is more fluid and coarsely granular, and contains various organs, chief amongst them in importance being the nucleus, which must be considered specially and may be put aside for the present . In considering the functions of ingestion and assimilation of food a distinction must be See also:drawn between those Protozoa which absorb solid food-particles, that is to say, which are holozoic in habit, and those which, being holophytic, saprophytic or parasitic in habit, absorb their nourishment in a See also:state of See also:solution . Only in holozoic forms is a special apparatus found for ingestion or digestion of food; in all other forms nutriment is absorbed by osmosis through the body-See also:wall, presumably at any point of the surface . In holozoic forms we must distinguish further those in which the protoplasm is naked at the surface from those in which the body is clothed by a firm cuticle or cell-membrane . In naked forms food-particles are taken in at any point of the body-surface, either by means of the pseudopodia, or by the action of flagella causing them to impinge upon the surface of the body . In either case the food is absorbed by the protoplasm simply flowing round it and engulfing it, and the food passes into the interior of the body in a tiny droplet of water forming what is termed a food-vacuole . Into the food-vacuole the surrounding protoplasm secretes digestive enzymes, so that each such vacuole represents a minute digestive cavity, in which the food is slowly digested, rendered soluble, and absorbed by the surrounding protoplasm . The insoluble See also:residue of the food is finally rejected by expelling the food-vacuole and its contents from the surface of the body at any convenient point . The simple process of food-absorption described above for the more primitive naked forms is necessarily modified in detail, though not in principle, in corticate Protozoa, that is to say, in forms provided with a cuticle . In the first place, it becomes necessary to have a special aperture for the ingestion of food, a cell-mouth or cytostome . Primitively the cytostome is a simple pore or interruption of the cuticle, but in forms more highly evolved the aperture is prolonged inwards in the form of a See also:tube lined by ectosarc and cuticle, forming a gullet or See also:oesophagus which ends in the endoplasm . Food-particles are forced by the action of cilia or flagella down the oesophagus and collect at the bottom of it in a droplet of water which, after reaching a certain size, passes into the endoplasm as a food-vacuole in which the food is digested . For rejection of the insoluble residue of the food-vacuoles, a special pore or cell-anus (cvtopyge) may be present . In the Ciliata there is often a distinct anal tube visible at all times, but as a rule the anus is only visible at the moment that faecal matter is being ejected from it, though fine sections show that the pore is a constant one . In the higher Flagellata, on the other hand, the oesophageal ingrowth forms commonly a sort of cloacal cavity, into which the contractile vacuole or vacuoles See also:discharge themselves, and into which also the food-vacuoles evacuate their residues . Besides the food-vacuoles already described, and the nuclear apparatus presently to be dealt with, the endoplasm may contain various metaplastic products, that is to say, bodies to be regarded as stages in the upward or downward metabolism of the proto-. plasmic substance . Such substances may take the form of coarse granules of various kinds, crystals, vacuoles or droplets of fatty or oily nature, pigment-grains, and other bodies . In the holophytic Flagellata the endoplasm contains also various organs proper to the vegetable cell, such as chlorophyll-bodies (chromatophores), pyrenoids, grains of a starchy nature (paramylum), and so forth, which need not be described here in detail . The nucleus in Protozoa is usually a compact, fairly conspicuous structure, composed of chromatin combined in various ways with an achromatic substance or substances . Sometimes the chromatin is distributed in smaller masses through the nucleus, producing a granular type of nucleus; more often the chromatin is more or less concentrated in a central mass forming a so-calledkaryosome, consisting of an achromatic plastinoid substance impregnated with chromatin . If the karyosome is large and there is very little chromatin between it and the nuclear membrane, the nucleus is of the type termed vesicular . A nuclear membrane is not, however, always present, and true nucleoli, of the type found in the nuclei of metazoan cells, are not found in Protozoa . A given individual may have more than one nucleus, and the number present may amount to many thousands, as in the plasmodia of See also:Mycetozoa . In such cases the nuclei may be all of one kind, that is to say, not markedly different in size, structure or function, so far as can be seen; or there may be a pronounced morphological differentiation of the nuclei correlated with a difference of function . Thus in the class Infusoria two nuclei are found in each individual; a macronucleus which is somatic in function, that is to say, which regulates the metabolism and vital processes of the body generally, and the micro-nucleus, which is generative in function, that is to say, which remains in reserve during the See also:ordinary, " vegetative " life of the organism and becomes active during the act of syngamy, after which the effete macronucleus is absorbed or See also:cast out and a new somatic nucleus is formed from portions of the micronuclei which have undergone fusion in the sexual act . Thus the micro-nucleus of the Infusoria can be compared in a general way with the germ-plasm of the Metazoa, like which it remains inactive until the sexual union . On the other hand, in some Flagellata a differentiation of the nucleus of quite a different type is seen, a smaller, kinetic nucleus being separated off from the larger, trophic or principal nucleus . The kinetic nucleus has the function, apparently, of controlling the locomotor apparatus, so that the specialization of these two nuclei is of a kind quite different from that seen in the Infusoria . Besides the nuclear substance which is concentrated to form the principal nucleus or nuclei, there may be present also extranuclear granules of chromatin, so-called chromidia, scatteredt throughout the whole or some part of the protoplasmic body . Chromidia may be normally present in addition to the principal nucleus, or may be formed from the principal nucleus during certain phases of the life-cycle . In some cases the entire nucleus may become resolved temporarily into chromidia, from which a new nucleus may be formed again later by condensation and concentration of the scattered granules . When the chromidia are numerous and closely packed they may form a so-called chromidial network (Chromidial-Netz) . Recent observations on the reproduction of some See also:Sarcodina have shown that the chromidia may possess great importance in the life-cycle as representing generative chromatin which, like the See also:micronucleus of the Infusoria mentioned above, remains in reserve until, by the process of syngamy, the nuclear apparatus is renewed;; while the principal nuclei represent, like the macronuclei,,, somatic or vegetative chromatin which becomes effete and is cast off or absorbed when syngamy takes place . These questions will be discussed further below . It was formerly supposed that the lowest Protozoa were entirely without a nucleus, and on this supposition E . Haeckel attempted to establish a class named by him Monera, defined as Protozoa consisting of protoplasm alone, in which a nucleus was not differentiated . To this class were referred various organisms whose alleged archaic nature was expressed by such names as See also:Protogenes primordialis, organisms which, like so many other of the primitive forms of animal life described by Haeckel, have been seen by that naturalist alone up to the present . In all Protozoa that have been examined by modern methods a nucleus in some form has been demonstrated to exist, and it must be supposed, until See also:proof to the contrary be forthcoming, that in the case of the so-called Monera either the nucleus was overlooked owing to defective technique, or it had been temporarily resolved into chromidia . The nuclear apparatus may be supplemented by other bodies of which the nature is not always clear . Such is the so-called " Nebenkern " of Paramoeba eilhardi, apparently of the nature of a centrosome . Sometimes the karyosome acts like a . centrosome during the division of the nucleus, and sometimes in a general way, but can be given special names in special cases true centrosomes are present . Flagella also commonly arise from basal granules of a centrosomic nature, blepharoplasts in the correct sense of the term;l these blepharoplasts are always in connexion with the nucleus, or with the kinetic nucleus if there is one distinct from the trophic nucleus, as in the genus Trypanosoma and allied forms . Reproduction of the Protozoa.—The mode of reproduction in these organisms is the same as that of the cell generally, and takes always the form of fission of some kind; that is to say, of division of the body into smaller portions, each of which represents a See also:young individual . The division of the body is preceded by that of the nucleus, if single, or of each nucleus in; the cases where there are two different nuclei; if, however, more than one nucleus of the same kind be present, the nuclei may be simply shared amongst the daughter-individuals, this mode of division being known as plasmotomy . Other organs of the body may either, like the nucleus, undergo fission, or may be formed afresh in the daughter-individuals . The division of the nucleus in Protozoa may take place by the See also:direct method or by means of mitosis . Direct division, without mitosis, is of very common occurrence; the division may be simple or multiple, that is to say, into only two parts, or into a number of fragments formed simultaneously . An extreme case of multiple fission is seen in the formation of the microgametes of Coccidium schubergi, where the nucleus breaks up into a great number of chromidia, which become concentrated in patches to form the several daughter-nuclei . In some cases, on the other hand, multiple daughter-nuclei are formed by rapidly repeated simple division of the See also:parent nucleus . The mode of division may be different in different nuclei of the same individual; thus in the Infusoria the macronucleus divides by direct division, the micronucleus by mitosis . The mitosis of the Protozoa is far from being of the See also:uniform stereotyped See also:pattern seen in the Metazoa, but, as might have been expected, often shows a much simpler and more primitive condition . Centrosomes are often absent, and their place may be taken, as stated above, by other bodies . The nuclear membrane may be retained throughout the mitosis . Definite chromosomes can, as a rule, be made out, but the chromosomes are often very numerous and minute, without definite form, and See also:divide irregularly . Much remains to be done in studying the mitosis of the Protozoa, but it is probable that wider knowledge will show many conditions intermediate between direct division and perfect mitosis . The simplest method of fission in Protozoa is that termed binary, where the body divides into two halves, which may be equal and similar, so that the result is two See also:sister-individuals impossible to distinguish as parent and offspring . In many cases of binary fission, however, the resulting daughter-individuals may be markedly unequal in size, so that one maybe distinguished as the parent, the other as the offspring . If the daughter-individual be relatively very small, and formed in a more or less imperfect condition at first, the process is termed gemmation or budding . The buds formed in this way may be either external, formed on the surface of the body, or internal, that is, formed in special internal cavities, from which the offspring are later set free, as in many Acinetaria . Gemmation may be correlated with multiple nuclear fission in such a way that buds are formed over the whole body surface of the organism, which thereby under-goes a process of simultaneous multiple fission into numerous daughter-individuals . Rapid multiple fission of this kind is termed sporulation, and is a form of reproduction which is of common occurrence, especially in parasitic forms . Usually, the central portion of the parent body remains over as a residual body (Restkorper), but sometimes the parent organism is entirely resolved into the daughter-individuals, which are termed spores ' The kinetic nucleus of Trypanosoma is sometimes, but in the writer's See also:opinion wrongly, named centrosome or blepharoplast; the bodies to which cytologists give these names are achromatic bodies; the kinetic nucleus is a true See also:chromatic nucleus . The question of the centrosome in Protozoa is discussed by R . See also:Goldschmidt and M . Popoff . (see GREGARINES, See also:COCCIDIA, &C.) . Life-cycles of the Protozoa.—It is probable that in all Protozoa, as in the Metazoa, the life-history takes its course in a See also:series of recurrent cycles of greater or less extent, a fixed_point, as it were, in the cycle being marked by the act of syngamy, or conjugation, which represents, apparently, a process for recuperation of the waning vital powers of the organism . It is true that in many types of Protozoa syngamy is not known as yet to occur, but in all species which have been thoroughly investigated syngamy in some form has been observed, and there is nothing to lead to the belief that the sexual process is not of universal occurrence in the Protozoa . The life-cycle of a given species may be very simple or it may be extremely complex, the organism occurring under many different forms at different phases or periods of its development . The polymorphism of the Protozoa is best considered under three categories, according to the three See also:main causes to which it is due, namely, first, polymorphism due to See also:adaptation to different conditions of existence; secondly, polymorphism due to See also:differences of size and structure during growth; thirdly, polymorphism due to the differentiation of individuals in connexion with the process of syngamy or sexual conjugation . x . Polymorphism in Relation to Life-conditions.—As a protection against unfavourable conditions, or for other reasons, most Protozoa have the See also:power of passing into a resting condition, during which the vital functions may be wholly or in part suspended . In the resting phase the animal usually becomes enveloped in a resistant membrane or cyst secreted by it, and is then said to be encysted . The formation of a cyst may be a response to conditions of various kinds . Very commonly it is formed to protect the organism against a See also:change of medium, as in the case of See also:freshwater forms liable to See also:desiccation, or of parasites about to pass out of the bodies of their hosts . In other cases the organism passes into the resting state in order 'to absorb ingested nutriment or in order to enter upon reproductive phases . As a preparation for encystment, organs of locomotion, if present, are retracted or cast off; contractile vacuoles cease to be formed; and the food-vacuoles disappear, usually by digestion of their contents and rejection of the waste residue . The body becomes rounded off and more or less spherical in form, and the protoplasm becomes denser, that is, less fluid and more opaque, but at the same See also:time of diminished specific gravity, by loss of water . The cyst is then secreted at the surface as a layer of varying thickness and toughness . In the encysted condition many Protozoa are capable of being transported by the wind_ a fact which explains their appearance in infusions and. quids exposed to the air . In favourable conditions the cysts germinate, that is to say, the envelope is dissolved and the contained See also:organ-ism or organisms are set free to enter upon the strenuous life once more . In the Mycetozoa, organisms adapted to a semi-terrestrial life in moist surroundings, the protoplasm is capable, when desiccated, of passing into a tough condition resembling sealing-See also:wax, which, when moistened, assumes again its normal appearance and active condition . Resting phases, analogous to encystment, are seen in the spores of various forms, especially those of parasitic habit, which are commonly enclosed in tough, resistant envelopes or sporocysts, and enveloped as a protection against change of medium or of host . Within the sporocyst multiplication of the sporoplasm may take place to form more or fewer sporozoites . The sporocysts usually show definite symmetry and structure, infinitely variable in different species . In a suitable medium the spores germinate by rupture of the sporocysts and See also:escape of the contents . 2 . Polymorphism in Relation to Growth and Development.—In many species of Protozoa there is hardly any difference to be observed between different individuals during their active phases except in size . Those individuals about to multiply by fission are slightly above the normal in dimensions: op the other hand, those resulting from recent fission will be smaller than the See also:average; and such differences are, it need hardly be said, more pronounced when the fission is of the unequal binary type, or in cases of gemmation or multiple fission . In cases also where a given See also:strain of a species is becoming senile, it is sometimes observed that the individuals are markedly undersized on the average . On the other hand, it is often the case that the young individuals resulting from a recent act of multiplication may differ from adult individuals of the species, not merely in size, but in structural characters, to such an extent that their relationship to the adult forms could not be determined by simple inspection without other See also:evidence . This is especially true of those species in which multiplication by sporulation occurs, giving rise to numerous small spores which may at first be in a resting condition, enveloped in protective sporocysts, but which sooner or later become free, motile individuals known technically as swarm-spores . Thus in many Sarcodina the adult is a large amoeboid organism which produces by sporulation a great number of relatively minute swarm-spores . These may be either, as in the common Amoeba See also:proteus, amoeboid organisms, so-called amoebulae or pseudopodiospores, or, as in the Foraminifera and Radiolaria, flagellated organisms, so-called flagellulae or flagellispores . Sometimes, as in many Mycetozoa, amoeboid and flagellated phases may succeed each See also:ether rapidly in the development of the swarm-spores . The See also:familiar Noctiluca miliaris is another instance of a species which produces by sporulation numerous tiny swarm-spores quite different from the parent form in their characters . Such instances could be multiplied indefinitely amongst the Protozoa . When the young individuals differ greatly from the adults in structure and appearance they may be regarded as larval forms, and it is interesting to See also:note that such forms appear to be just as much recapitulative, in the phylogenetic sense, as are the larvae of many Metazoa . A striking instance is that of the Acinetaria, in which the swarm-spores produced by gemmation are ciliated, and thus betray See also:affinities with the Ciliata which could hardly be suspected from a study of the adult forms alone . Similarly, in the genus Trypanosoma, the young forms often show a Herpetomonas-like structure which is probably of phyletic significance . The swarm-spores of Sarcodina and of Noctiluca mentioned above can, perhaps, be regarded in the same light . On the other hand, many larval forms cannot be considered as exhibiting recapitulative characters, but merely as adaptations to environment or other special life-conditions . This is especially true, as in Metazoa, of parasitic forms, subject as they are to great vicissitudes, to See also:cope with which the most finely adjusted adaptations are necessary on the part of the organism . 3 . Polymorphism in Relation to See also:Sex.—In all Protozoa of which the life-cycle has been made known in its entire course, a process of syngamy or sexual union has been found to occur . There are still many forms in which syngamy remains to be discovered: this is true even of some See also:groups of considerable extent . It is quite possible, therefore, that Protozoa exist in which syngamy does not occur . In view, however, of the widespread occurrence of sexual processes amongst unicellular organisms, both of animal and vegetable nature, and the fact that extended observation continually brings to light new instances of this kind, it is safer, in cases amongst the Protozoa in which syngamy is not known to occur, to explain its apparent absence by the imperfections of the present state of our know-ledge, than to suppose that in such forms sexual phenomena are entirely lacking in the life-cycle.' The process of syngamy, though greatly diversified in different forms, consists essentially of one and the same process in all cases; namely, the fusion of nuclear matter from two distinct individuals . Plus ca change, plus c'est la mgme See also: |