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Monas St.; Anthophysa Bory (Fig. 37, 13).
(b) One flagellum, usually the
longer, turned backwards Bodonidae
Bodo St. (Fig. 38).
C. Flagella 2, equal and similar Amphimonadidae
Amphimonas Duj.; Diplomita K. (Fig. 37, 10);
Rhipidodendron St. (Fig. 37, 14).
D. Flagella 3 Trimastigidae
Dallingeria K. (Fig. 37, 6); Costia Leclercq.
E. Flagella 4 or more: mostly parasitic
in Metazoa Polymastigidae
Trichomonas Donne; Tetramitus Perty (Fig. 37, 7);
Hexamitus Duj.; Lamblia Blanchard.
F. Flagella numerous, sometimes
constituting a complete ciliiform
investment, and occasionally
accompanied by an undulating
membrane: parasitic in Metazoa.
(a) Flagella long: nucleus single:
parasitic in insects Trichonymphidae
Dinenympha Leidy; Joenia Grassi; Pyrsonympha
Leidy; Trichonympha Leidy; Lophomonas St.;
Maupasia Schew.
(b) Flagella short, ciliiform,
uniformly distributed: nuclei
very numerous, all similar:
parasitic in Amphibia Opalinidae
Opalina Purkinje and Valentin (Fig. 41).
Volvocaceae
A. Cells usually isolated, separating after
fission or brood-formation. Usually
green (sometimes red), more rarely
colourless saprophytes Chlamydomonadidae
Chlamydomonas Ehrb.; Phacotus Perty; Polytoma Ehrb.;
Sphaerella Sommerf. (Fig. 43); Zoochlorella.
B. Cells multiplying in the active state by
radial divisions in the same plane
and usually incurving to form a
spherical colony, united in a
gelatinous investment, sometimes
traversed by plasmic threads Volvocidae
Gonium O.F.M.; Eudorina Ehrb.; Pandorina Bory (Fig. 45);
Stephanosphaera Cohn; Volvox L. (Fig. 44).
Fig. 37.—Various forms of Flagellata. 2, 6-8, 10, 13, 14, Protomastigaceae; 11,
12, Chrysomonadaceae; 9, Cryptomonadaceae; 1, 3, Euglenaceae; 4,
Pantostomata: note branched stalk in 13; branched tubular theca in 14;
distinct thecae in 11; stalk and theca in 10. In 2, flagellate (a) and amoeboid
(b) phases are shown; in 5, flagellate (a) and Heliozoan (b) phases[116]; in 8
are shown two stages in the ingestion of a food particle (f); chr, plastoids;
c.vac, contractile vacuole; f, food particle; g, gullet; l, theca; nu, nucleus; p,
protoplasm; per, peristome; v.i, vacuole of ingestion. (From Parker and
Haswell, mostly from Bütschli's Protozoa.)
The modes of nutrition are threefold: the simplest forms live in liquids
containing decaying organic matter which they absorb through their
surface ("saprophytic"): others take in food either Amoeba fashion,
or into a vacuole formed for the purpose, or into a definite mouth
("holozoic"): others again have coloured plastids, green or brown or
yellow ("holophytic"), having the plant's faculty of manufacturing their
own food-supply. But we meet with species that show
chromatophores at one time and lack them at another; or, again, the
same individual (Euglena) may pass from holozoic life to saprophytic
(Paramoeba, some Dinoflagellates) as conditions alter.
The flagellum has been shown by Fischer to have one of two forms:
either it is whip-like, the stick, alone visible in the fresh specimen,
being seen when stained to be continued into a long lash, hitherto
invisible; or the whole length is fringed with fine ciliiform lateral
outgrowths. If single it is almost always protruded as a tugging organ
("tractellum");[120] the chief exceptions are the Craspedomonads,
where it is posterior and acts as a scull ("pulsellum"), and some
Dinoflagellates, where it is reversible in action or posterior. In
addition to the anterior flagellum there may be one or more posterior
ones, which trail behind as sense organs, or may anchor the cell by
their tips. Dallingeria has two of these, and Bodo saltans a single
anterior anchoring lash, by which they spring up and down against
the organic débris among which they live, and disintegrate it. The
numerous similar long flagella of the Trichonymphidae afford a
transition in the genus Pyrsonympha to the short abundant cilia of
Opalina, usually referred to the Ciliate Infusoria.
In some cases the flagellum (or flagella) is inserted into a definite pit,
which in allied forms is the mouth-opening. The contractile vacuole is
present in the fresh-water forms, but not in all the marine ones, nor
in the endoparasites. It may be single or surrounded by a ring of
minute "formative" vacuoles or discharge into a permanently visible
"reservoir." This again may discharge directly to the surface or
through the pit or canal in which the flagellum takes origin (Euglena).
As we noted above (p. 40), the study of the Flagellates has been
largely in the hands of botanists. After the work of Bütschli in Bronn's
Thier-Reich, Klebs[126] took up their study; and the principal
monographs during the last decade have appeared in Engler and
Prantl's Pflanzenfamilien, where Senn[127] treats the Flagellates
generally, Wille[128] the Volvocaceae, and Schütt the "Peridiniales" or
Dinoflagellata;[129] while only the Cystoflagellata, with but two
genera, have been left to the undisputed sway of the zoologists.[130]
The human Tick fever of the Western United States and the epizootic
Texas fever are known to be due to blood parasites of the genus
Piroplasma (Babesia), of which the free state is that of a
Trypanosome. It appears certain that Texas fever, though due to Tick
bites, is not transferred directly from one beast to another by the
same Tick; but the offspring of a female Tick that has sucked an
infected ox contains Trypanosome germs, and will by their bites
infect other animals. It would seem probable that the virulence of the
Persian Tick (Argas persica) is due to similar causes. The Indian
maladies known as "Kala Azar" and "Oriental Sore" are
characterised by blood parasites, at first called after their discoverer
the "Leishman bodies," which have proved to be the effects of a
Piroplasma.
The divisions are only in two planes at right angles, so that the
young colony is at first a plate, but as the cells multiply the plate
bends up (as in the gastrulation of the double cellular plate of the
Nematode Cucullanus, Vol. II. p. 136), and finally forms a hollow
sphere bounded by a single layer of cells: the site of the original
orifice may be traced even in the adult as a blank space larger than
exists elsewhere. Among the cells of the young colony some cease
to divide, but continue to grow at an early period, and these are
destined to become in turn the mothers ("parthenogonidia") of a new
colony; they begin segmenting before the colony of which they are
cells is freed. The young colonies are ultimately liberated by the
rupture of the sphere as small-sized spheres, which henceforth only
grow by enlargement of the sphere as a whole, and the wider
separation of the vegetative cells. Thus the vegetative cells soon
cease to grow; all the supply of food material due to their living
activities goes to the nourishment of the parthenogonidia, or the
young colonies, as the case may be. These vegetative cells have
therefore surrendered the power of fission elsewhere inherent in the
Protist cell. Moreover, when the sphere ruptures for the liberation of
the young colonies, it sinks and is doomed to death, whether
because its light-loving cells are submerged in the ooze of the
bottom, or because they have no further capacity for life. When
conjugation is about to take place, it is the cells that otherwise would
be parthenogonidia that either act as oospheres or divide as
"spermogonia" to form a flat brood of minute yellow male cells
("sperms"). These resemble vegetative cells, in the possession of an
eye-spot and two contractile vacuoles, but differ in the enormously
enlarged nucleus which determines a beaked process in front. After
one of these has fused with the female cell ("oosphere") the product
("oosperm") encysts, passes into a stage of profound rest, and finally
gives rise to a new colony. The oospheres and sperm-broods may
arise in the same colony or in distinct ones, according to the species.
Stephanosphaera has its eight cells spindle shaped, and lying along
equidistant meridians of its sphere; in vegetative reproduction each
of these breaks up in its place to form a young colony, and the eight
daughter-colonies are then freed. In conjugation, each cell of the
colony breaks up into broods of 4, 8, 16, or 32 small gametes, which
swim about within the general envelope, and pair and fuse two and
two: this is "isogamous," "endogamous" conjugation. In Pandorina
(Fig. 45) the cells are rounded, and are from 16 to 32 in each colony.
The vegetative reproduction in this, as in Eudorina, is essentially the
same as in Stephanosphaera. In conjugation the cells are set free,
and are of three sizes in different colonies, small (S), medium (M),
and large (L). The following fusions may occur: S × S, S × M, S × L,
M × M, M × L. Thus the large are always female, as it were, the
medium may play the part of male to the large, female to the small;
the small are males to the medium and to the large. The medium
and small are capable, each with its like, of equal, undifferentiated
conjugation; so that we have a differentiation of sex far other than
that of ordinary, binary sex. Eudorina, however, has attained to
"binary sex," for the female cells are the ordinary vegetative cells, at
most a little enlarged, and the male cells are formed by ordinary cells
producing a large flat colony of sixty-four minute males or sperms. In
some cases four cells at the apex of a colony are spermogonia,
producing each a brood of sperms, while the rest are the oospheres.
The transition to Volvox must have arisen through the sterilisation of
the majority of cells of a colony for the better nutrition of the few that
are destined alone for reproduction.