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 Fig. 20.—Various forms of Heliozoa. In 3, a is the entire animal and b the
flagellula; c.vac, contractile vacuole; g, gelatinous investment; nu, nucleus;
psd, pseudopodia; sk, siliceous skeleton; sp, spicules. (From Parker and
Haswell, after other authors.)

Each of these divides, and the two sister cells then conjugate after
the same fashion as in Actinophrys, but the nuclear divisions to form
the coupling nucleus are two in number, i.e. the nucleus divides into
two, one of which goes to the surface as the first polar body, and the
sister of this again divides to form a second polar body (which also
passes to the surface) and a pairing nucleus.[84] The two cells then
fuse completely, and surround themselves with a second gelatinous
cyst wall, separated from the outer one by a layer of siliceous
spicules. The nucleus appears to divide at least twice before the
young creep out, to divide immediately into as many Actinophrys-like
cells as there were nuclei; then each of these multiplies its nuclei, to
become apocytial like the adult form.
 Fig. 21.—Diagram illustrating the conjugation of Actinosphaerium. 1, Original
cell; 2, nucleus divides to form two, N2N2; 3, each nucleus again divides to
form two, N3 and n3, the latter passing out with a little cytoplasm as an
abortive cell; 4, repetition of the same process as in 3; 5, the two nuclei N4
have fused in syngamy to form the zygote nucleus Nz.

Schaudinn admits 24 genera (and 7 doubtful) and 41 species (and

18 doubtful). None are known fossil. Their geographical distribution
is cosmopolitan, as is the case with most of the minute fresh-water
Protista; 8 genera are exclusively marine, and Orbulinella has only
been found in a salt-pond; Actinophrys sol is both fresh-water and
marine, and Actinolophus has 1 species fresh-water, the other
marine. One of the 14 species of Acanthocystis is marine; the
remaining genera and species are all inhabitants of fresh water.[85]

4. Radiolaria
Sarcodina with the protoplasm divided by a perforated chitinous
central capsule into a central mass surrounding the nucleus, and an
outer layer; the pseudopodia radiate, never anastomosing enough to
form a marked network; skeleton either siliceous, of spicules, or
perforated; or of definitely arranged spicules of proteid matter
(acanthin), sometimes also coalescing into a latticed shell;
reproduction by fission and by zoospores formed in the central
capsule. Habitat marine, suspended at the surface (plankton), at
varying depths (zonarial), or near the bottom (abyssal).

Fig. 22.—Collozoum inerme. A, B, C, three forms of colony; D, small colony with

central capsules (c.caps), containing nuclei, and alveoli (vac) in ectoplasm;
 E, isospores, with crystals (c); F, anisospores; nu, nucleus. (From Parker
and Haswell.)

The following is Haeckel's classification of the Radiolaria:—

I. Porulosa (Holotrypasta).—Homaxonic, or nearly so. Central capsule
spherical in the first instance; pores numerous, minute, scattered; mostly
pelagic.
A. Spumellaria (Peripylaea).—Pores evenly scattered; skeleton of solid
siliceous spicules, or continuous, and reticulate or latticed, rarely absent;
nucleus dividing late, as an antecedent to reproduction.

B. Acantharia (Actipylaea).—Pores aggregated into distinct areas;

skeleton of usually 20 centrogenous, regularly radiating spines of acanthin,
whose branches may coalesce into a latticed shell; nucleus dividing early.

II. Osculosa (Monotrypasta).—Monaxonic; pores of central capsule limited

to the basal area (osculum), sometimes accompanied by two (or more)
smaller oscula at apical pole, mostly zonarial or abyssal.
C. Nassellaria (Monopylaea).—Central capsule ovoid, of a single layer;
pores numerous on the operculum or basal field; skeleton siliceous, usually
with a principal tripod or calthrop-shaped spicule passing, by branching,
into a complex ring or a latticed bell-shaped shell; nucleus eccentric, near
apical pole.

D. Phaeodaria (Cannopylaea, Haeck.; Tripylaea, Hertw.).—Central

capsule spheroidal, of two layers, in its outer layer an operculum, with
radiate ribs and a single aperture, beyond which protrudes the outer layer;
osculum basal, a dependent tube (proboscis); accessory oscula, when
present, simpler, usually two placed symmetrically about the apical pole;
skeleton siliceous, with a combination of organic matter, often of hollow
spicules; nucleus sphaeroidal, eccentric; extracapsular protoplasm
containing an accumulation of dusky pigment granules ("phaeodium").
 Fig. 23.—Actinomma asteracanthion. A, the shell with portions of the two outer
spheres broken away; B, section showing the relations of the skeleton to
the animal, cent.caps, Central capsule; ex.caps.pr, extra-capsular
protoplasm: nu, nucleus; sk.1, outer, sk.2, middle, sk.3, inner sphere of
skeleton. (From Parker and Haswell, after Haeckel and Hertwig.)

A. Spumellaria.

Sublegion (1). Collodaria.[86]—Skeleton absent or of detached spicules;

colonial or simple.
Order i. Colloidea.—Skeleton absent. (Families 1, 2.) Thalassicolla Huxl.;
Thalassophysa Haeck.; Collozoum Haeck.; Collosphaera J. Müll.; Actissa
Haeck.

Order ii. Beloidea.—Skeleton spicular. (Families 3, 4.)

Sublegion (2). Sphaerellaria.—Skeleton continuous, latticed or spongy,

reticulate.
Order iii. Sphaeroidea.—Skeleton of one or several concentric spherical
shells; sometimes colonial. (Families 5-10.) Haliomma Ehrb.; Actinomma
Haeck. (Fig. 23).

Order iv. Prunoidea.—Skeleton a prolate sphaeroid or cylinder, sometimes

constricted towards the middle, single or concentric. (Families 11-17.)

Order v. Discoidea.—Shell flattened, of circular plan, simple or concentric,

rarely spiral. (Families 18-23.)

Order vi. Larcoidea.—Shell ellipsoidal, with all three axes unequal or

irregular, sometimes becoming spiral. (Families 24-32.)[87]
 Fig. 24.—Xiphacantha (Acantharia). From the surface. The skeleton only, × 100,
(From Wyville Thomson.)

B. Acantharia.

Order vii. Actinelida.—Radial spines numerous, more than 20, usually

grouped irregularly. (Families 33-35.) Xiphacantha Haeck.

Order viii. Acanthonida.—Radial spines equal. (Families 36-38.)

Order ix. Sphaerophracta.—Radial spines 20, with a latticed spherical

shell, independent of, or formed from the reticulations of the spines.
(Families 39-41.) Dorataspis Haeck. (Fig. 25, A).

Order x. Prunophracta.—Radial spines 20, unequal; latticed shell,

ellipsoidal, lenticular, or doubly conical. (Families 42-44.)

C. Nassellaria.

Order xi. Nassoidea.—Skeleton absent. (Family 45.)

Order xii. Plectoidea.—Skeleton of a single branching spicule, the

branches sometimes reticulate, but never forming a latticed shell or a
sagittal ring. (Families 46-47.)

Order xiii. Stephoidea.—Skeleton with a sagittal ring continuous with the

branched spicule, and sometimes other rings or branches. (Families 48-
 51.) Lithocercus Théel (Fig. 26, A).

Order xiv. Spyroidea.—Skeleton with a latticed shell developed around the

sagittal ring (cephalis), and constricted in the sagittal plane, with a lower
chamber (thorax) sometimes added. (Families 52-55.)

Order xv. Botryoidea.—As in Spyroidea, but with the cephalis 3-4 lobed;
lower chambers, one or several successively formed. (Families 56-58.)

Order xvi. Cyrtoidea.—Shell as in the preceding orders, but without lobing

or constrictions. (Families 59-70.) Theoconus Haeck. (Fig. 25, B).

D. Phaeodaria.

Order xvii. Phaeocystina.—Skeleton 0 or of distinct spicules; capsule

centric. (Families 71-73.) Aulactinium Haeck. (Fig. 26, B).

Order xviii. Phaeosphaeria.—Skeleton a simple or latticed sphere, with no

oral opening (pylome); capsule central. (Families 74-77.)

Order xix. Phaeogromia.—Skeleton a simple latticed shell with a pylome

at one end of the principal axis; capsule excentric, sub-apical. (Families 78-
82.) Pharyngella Haeck.; Tuscarora Murr.; Haeckeliana Murr. (Fig. 28).

Order xx. Phaeoconchia.—Shell of two valves, opening in the plane

("frontal") of the three openings of the capsule. (Families 83-85.)

We exclude Haeckel's Dictyochida, with a skeleton recalling that of

the Stephoidea, but of the impure hollow substance of the
Phaeodaria (p. 84). They rank now as Silicoflagellates (p. 114).

The Radiolarian is distinguished from all other Protozoa by the

chitinous central capsule, so that its cytoplasm is separated into an
outer layer, the extracapsular protoplasm (ectoplasm), and a central
mass, the intracapsular, containing the nucleus.[88]
The extracapsular layer forms in its substance a gelatinous mass, of
variable reaction, through which the plasma itself ramifies as a
network of threads ("sarcodictyum"), uniting at the surface to
constitute the foundation for the pseudopodia. This gelatinous matter
constitutes the "calymma." It is largely vacuolated, the vacuoles
("alveoli"), of exceptional size, lying in the nodes of the plasmic
network, and containing a liquid probably of lower specific gravity
than seawater; and they are especially abundant towards the
surface, where they touch and become polygonal. On mechanical
irritation they disappear, to be formed anew after an interval, a fact
that may explain the sinking from the surface in disturbed water. This
layer may contain minute pigment granules, but the droplets of oil
and of albuminous matter frequent in the central layer are rare here.
The "yellow cells" of a symbiotic Flagellate or Alga, Zooxanthella, are
embedded in the jelly of all except Phaeodaria, and the whole
ectosarc has the average consistency of a firm jelly.

The pseudopodia are long and radiating, with a granular external

layer, whose streaming movements are continuous with those of the
inner network. In the Acantharia they contain a firm axial filament,
like that of the Heliozoa, which is traceable to the central capsule;
and occasionally a bundle of pseudopodia may coalesce to form a
stout process like a flagellum ("sarcoflagellum"). Here, too, each
spine, at its exit from the jelly, is surrounded by a little cone of
contractile filaments, the myophrisks, whose action seems to be to
pull up the jelly and increase the volume of the spherical body so as
to diminish its density.

Fig. 25.—Skeletons of Radiolaria. A, Dorataspis; B, Theoconus. (After Haeckel.)

The intracapsular protoplasm is free from Zooxanthella except in the
Acantharia. It is less abundantly vacuolated, and is finely granular. In
the Porulosa it shows a radial arrangement, with pyramidal stretches
of hyaline plasma separated by intervals rich in granules. Besides
the alveoli with watery contents, others are present with albuminoid
matter in solution. Oil-drops, often brilliantly coloured, occur either in
the plasma or floating in either kind of vacuole; and they are often
luminous at night. Added to these, the intracapsular plasm contains
pigment-granules, most frequently red or orange, passing into yellow
or brown, though violet, blue, and green also occur. The
"phaeodium,"[89] however, that gives its name to the Phaeodaria, is
an aggregate of dark grey, green, or brown granules which are
probably formed in the endoplasm, but accumulate in the
extracapsular plasm of the oral side of the central capsule. Inorganic
concretions and crystals are also found in the contents of the central
capsule, as well as aggregates of unknown composition, resembling
starch-grains in structure.

In the Monopylaea, or Nassellaria (Figs. 25, B, 26, A), the

endoplasm is differentiated above the perforated area of the central
capsule into a cone of radiating filaments termed the "porocone,"
which may be channels for the communication between the
exoplasm and the endoplasm, or perhaps serve, as Haeckel
suggests, to raise, by their contraction, the perforated area: he
compares them to the myophane striae of Infusoria. In the
Phaeodaria (Fig. 26, B), a radiating laminated cone is seen in the
outermost layer of the endoplasm above the principal opening
("astropyle"), and a fibrillar one around the two accessory ones
("parapyles"); and in some cases, continuous with these, the whole
outer layer of the endoplasm shows a meridional striation.

The nucleus is contained in the endoplasm, and is always at first

single, though it may divide again and again. The nuclear wall is a
firm membrane, sometimes finely porous. If there are concentric
shells it at first occupies the innermost, which it may actually come to
enclose, protruding lobes which grow through the several
perforations of the lattice-work, finally coalescing outside completely,
so as to show no signs of the joins. In the Nassellaria a similar
process usually results in the formation of a lobed nucleus,
contained in an equally lobed central capsule. The chromatin of the
nucleus may be concentrated into a central mass, or distributed into
several "nucleoli," or it may assume the form of a twisted, gut-like
filament, or, again, the nuclear plasm may be reticulated, with the
chromatin deposited at the nodes of the network.

Fig. 26.—A, Lithocercus annularis, with sagittal ring (from Parker and Haswell).
B, Aulactinium actinastrum. C, calymma; cent.caps., km, central capsule;
Ext.caps.pr., Extracapsular, and Int.caps.pr., intracapsular protoplasm; n,
nu, nucleus; op, operculum; ph, phaeodium; psd, pseudopodium; Skel.,
skeleton; z, Zooxanthella. (From Lang's Comparative Anatomy, after
Haeckel.)

The skeleton of this group varies, as shown in our conspectus, in the

several divisions.[90] The Acantharia (Figs. 24, 25, A) have a
skeleton of radiating spines meeting in the centre of figure of the
endoplasm, and forcing the nucleus to one side. The spines are
typically 20 in number, and emerge from the surface of the regular
spherical forms (from which the others may be readily derived)
radially, in five sets of four in the regions corresponding to the
equator and the tropics and polar circles of our world. The four rays
of adjacent circles alternate, so that the "polar" and "equatorial" rays
are on one set of meridians 90° apart, and the "tropical" spines are
on the intermediate meridians, as shown in the figures. By tangential
branching, and the meeting or coalescence of the branches,
reticulate (Figs. 23, 24, 25) and latticed shells are formed in some
families, with circles of openings or pylomes round the bases of the
spines. In the Sphaerocapsidae the spines are absent, but their
original sites are inferred from the 20 circles of pylomes.

In the Spumellaria the simplest form of the (siliceous) skeleton is that

of detached spicules, simple or complex, or passing into a latticed
shell, often with one or more larger openings (pylomes). Radiating
spines often traverse the whole of the cavity, becoming continuous
with its latticed wall, and bind firmly the successive zones when
present (Fig. 23).

Calcaromma calcarea was described by Wyville Thomson as having

a shell of apposed calcareous discs, and Myxobrachia, by Haeckel,
as having collections of the calcareous Coccoliths and
Coccospheres. In both cases we have to do with a Radiolarian not
possessing a skeleton, but retaining the undigested shells of its food,
in the former case (Actissa) in a continuous layer, in the latter
(Thalassicolla) in accumulations that, by their weight, droop and pull
out the lower hemisphere into distinct arms.

The (siliceous) skeleton of the Nassellaria is absent only in the

Nassoidea, and is never represented by distinct spicules. Its simplest
form is a "tripod" with the legs downward, and the central capsule
resting on its apex. The addition of a fourth limb converts the tripod
into a "calthrop," the central capsule in this case resting between the
upturned leg and two of the lower three regarded as the
"anterolateral"; the odd lower leg, like the upturned one, being
"posterior." Again, the skeleton may present a "sagittal ring," often
branched and spiny (Fig. 26, A), or combined with the tripod or
calthrop, or complicated by the addition of one or more horizontal
rings. Another type is presented by the "latticed chamber"
surrounding the central capsule, with a wide mouth ("pylome") below.
This is termed the "cephalis"; it may be combined in various ways
with the sagittal ring and the tripod or calthrop; and, again, it may be
prolonged by the addition of one, two, or three chambers below, the
last one opening by a pylome (Fig. 25, B). These are termed
"thorax," "abdomen," and "post-abdomen" respectively.

In the Phaeodaria the skeleton may be absent, spicular (of loose or

connected spicules) or latticed, continuous or bivalve. It is composed
of silica combined with organic matter, so that it chars when heated,
is more readily dissolved, and is not preserved in fossilisation. The
spicules or lattice-work are hollow, often with a central filament
running in the centre of the gelatinous contents. The latticed
structure of the shell of the Challengeridae (Fig. 28) is so fine as to
recall that of the Diatomaceae. In the Phaeoconchida the shell is in
two halves, parted along the "frontal" plane of the three apertures of
the capsule.

Fig. 27.—Scheme of various possible skeletal forms deposited in the meshes of

an alveolar system, most of which are realised in the Radiolaria. (From
Verworn, after Dreyer.)

The central capsule (rarely inconspicuous and difficult, if not

impossible to demonstrate) is of a substance which resembles chitin,
though its chemical reactions have not been fully studied hitherto,
and indeed vary from species to species. It is composed of a single
layer, except in Phaeodaria, where it is double. The operculum in this
group, i.e. the area around the aperture, is composed of an outer
layer, which is radially thickened, and a thin inner layer; the former is
produced into the projecting tube ("proboscis").

Reproduction in the Radiolaria may be simple fission due to the

binary fission of the nucleus, the capsule, and the ectoplasm in
succession. If this last feature is omitted we have a colonial
organism, composed of the common ectoplasm containing
numerous central capsules; and the genera in which this occurs, all
belonging to the Peripylaea, were formerly separated (as
Polycyttaria) from the remaining Radiolaria (Monocyttaria). They may
either lack a skeleton (Collozoidae, Fig. 22), or have a skeleton of
detached spicules (Sphaerozoidae), or possess latticed shells
(Collosphaeridae) one for each capsule, and would seem therefore
to belong, as only differentiated by their colonial habit, to the several
groups having these respective characters. Fission has been well
studied in Aulacantha (a Phaeodarian) by Borgert.[91] He finds that in
this case the skeleton is divided between the daughter-cells, and the
missing part is regenerated. In cases where this is impossible one of
the daughter-cells retains the old skeleton, and the other escapes as
a bud to form a new skeleton.

Fig. 28.—Shells of Challengeridae: A, Tuscarora; B, Pharyngella; C,

Haeckeliana. (From Wyville Thomson.)

Two modes of reproduction by flagellate zoospores have been

described (Fig. 22). In the one mode all the zoospores are alike—
isospores—and frequently contain a crystal of proteid nature as well
as oil-globules. In the Polycyttaria alone has the second mode of
spore-formation been seen, and that in the same species in which
the formation of isospores occurs. Here "anisospores" are formed,
namely, large "mega-," and small "micro-zoospores." They probably
conjugate as male and female respectively; but neither has the
process been observed, nor has any product of such conjugation
(zygote) been recognised. In every case the formation of the
zoospores only involves the endoplasm: the nucleus first undergoes
brood division, and the plasma within the capsule becomes
concentrated about its offspring, and segregates into the spores; the
extracapsular plasm disintegrates.[92]

The Yellow Cells (Zooxanthella), so frequently found in the

Radiolaria were long thought to be constituents of their body.
Cienkowsky found that when the host died from being kept in
unchanged water, the yellow cells survived and multiplied freely,
often escaping from the gelatinised cell-wall as biflagellate
zoospores. The cell-wall is of cellulose. The cell contains two
chloroplastids, or plates coloured with the vegetal pigment
"diatomin." Besides ordinary transverse fission in the ordinary
encysted state in the ectoplasm of the host, when free they may
pass into what is known as a "Palmella-state," the cell-walls
gelatinising; in this condition they multiply freely, and constitute a
jelly in which the individual cells are seen as rounded bodies. They
contain starch in two forms—large hollow granules, not doubly
refractive, and small solid granules which polarise light. We may
regard them as Chrysomonadaceae (p. 113). Similar organisms
occur in many Anthozoa (see pp. 261, 339, 373 f., 396).
Diatomaceae (yellow Algae with silicified cell-walls) sometimes live
in the jelly of certain Collosphaera. Both these forms live in the state
known as "symbiosis" with their host; i.e. they are in mutually helpful
association, the Radiolarian absorbing salts from the water for the
nutrition of both, and the Alga or Flagellate taking up the CO2 due to
the respiration of the host, and building up organic material, the
surplus of which is doubtless utilised, at least in part, for the nutrition
of the host. A similar union between a Fungus and a coloured
vegetal ("holophytic") organism is known as a Lichen.

The Suctorian Infusorian Amoebophrya is parasitic in the ectoplasm

of certain Acantharia, and in the peculiar genus Sticholonche which
appears to be intermediate between this group and Heliozoa.

The Silicoflagellate family Dictyochidae are found temporarily

embedded in the ectoplasm of some of the Phaeocystina, and have
a skeleton of similar nature. Their true nature was shown by Borgert.

The Amphipod crustacean Hyperia[93] may enter the jelly of the

colonial forms, and feed there at will on the host.[94]

Haeckel, in his Monograph of the Radiolaria of the Challenger

enumerated 739 genera, comprising 4318 species; and Dreyer has
added 6 new genera, comprising 39 species, besides 7 belonging to
known genera. Possibly, as we shall see, many of the species may
be mere states of growth, for it is impossible to study the life-
histories of this group; on the other hand, it is pretty certain that new
forms are likely to be discovered and described. The Radiolaria are
found living at all depths in the sea, by the superficial or deep tow-
net; and some appear to live near the bottom, where the durable
forms of the whole range also settle and accumulate. They thus form
what is known as Radiolarian ooze, which is distinguished from other
shallower deposits chiefly through the disappearance by solution of
all calcareous skeletons, as they slowly fell through the waters
whereon they originally floated at the same time with the siliceous
remains of the Radiolaria. The greatest wealth of forms is found in
tropical seas, though in some places in cold regions large numbers
of individuals of a limited range of species have been found.

Radiolaria of the groups with a pure siliceous skeleton can alone be

fossilised, even the impure siliceous skeleton of the Phaeodaria
readily dissolving in the depths at which they live: they have been
generally described by Ehrenberg's name Polycystineae. Tripolis
(Kieselguhr) of Tertiary ages have been found in many parts of the
globe, consisting largely or mainly of Radiolaria, and representing a
Radiolarian ooze. That of the Miocene of Barbados contains at least
400 species; that of Gruppe at least 130. In Secondary and
Palaeozoic rocks such oozes pass into Radiolarian quartzites (some
as recent as the Jurassic). They occur also in fossilised excrement
(coprolites), and in flint or chert concretions, as far down as the
lowest fossiliferous rocks, the Cambrian. The older forms are simple
Sphaerellaria and Nassellaria. From a synopsis of the history of the
order in Haeckel's Monograph (pp. clxxxvi.-clxxxviii.) we learn that
while a large number of skeletal forms had been described by
Ehrenberg, Huxley in 1851 published the first account of the living
animal. Since then our knowledge has been extended by the labours
of Haeckel, Cienkowsky, R. Hertwig, Karl Brandt, and A. Borgert.

5. Proteomyxa
Sarcodina without a clear ectoplasm, whose active forms are
amoeboid or flagellate, or pass from the latter form to the former;
multiplying chiefly, if not exclusively, by brood-formation in a cyst. No
complete cell-pairing (syngamy) known, though the cytoplasms may
unite into plasmodia; pseudopodia of the amoeboid forms usually
radiate or filose, but without axial filaments. Saprophytic or parasitic
in living animals or plants.

This group is a sort of lumber-room for forms which it is hard to place

under Rhizopoda or Flagellata, and which produce simple cysts for
reproduction, not fructifications like the Mycetozoa. The cyst may be
formed for protection under drought ("hypnocyst"), or as a
preliminary to spore-formation ("sporocyst"). The latter may have a
simple wall (simple sporocyst), or else two or three formed in
succession ("resting cyst"), so as to enable it to resist prolonged
desiccation, etc.: both differing from the hypnocyst in that their
contents undergo brood formation. On encystment any indigestible
food materials are extruded into the cyst, and in the "resting cysts,"
which are usually of at least two layers, this faecal mass lies in the
space between them. The brood-cells escape, either as flagellate-
cells, resembling the simpler Protomastigina, called "flagellulae," and
which often become amoeboid (Fig. 29); or already furnished with
pseudopodia, and called "amoebulae," though they usually recall
Actinophrys rather than Amoeba. In Vampyrella and some others the
amoebulae fuse, and so attain a greater size, which is most probably
advantageous for feeding purposes. But usually it is as a uninucleate
cell that the being encysts. They may feed either by ingestion by the
pseudopodia, by the whole surface contained in a living host-cell, or
by passing a pseudopodium into a host-cell (Fig. 29 5). They may be
divided as follows:—

A. Myxoidea.—Flagella 1-3; zoospores separating at once.

1. Zoosporeae.—Brood-cells escaping as flagellulae, even if they become
amoeboid later. Ciliophrys Cienk.; Pseudospora Cienk. (Fig. 29).

2. Azoosporeae.—Cells never flagellate. Protomyxa Haeckel;

Plasmodiophora Woronin; Vampyrella Cienk.; Serumsporidium L. Pfeiffer.

B. Catallacta.—Brood-cells of cyst on liberation adhering at the centre to

form a spherical colony, multiflagellate; afterwards separating, and becoming
amoeboid. Magosphaera Haeckel (marine).[95]

Fig. 29.—Pseudospora lindstedtii. 1, 2, Flagellate zoospores; 3, young

amoebula, with two contractile vacuoles, one being reconstituted by three
minute formative vacuoles; 4, 5, an amoebula migrating to a fungus hypha
through the wall of which it has sent a long pseudopodium; 6, amoebula full-
grown; 7, 8, mature cells rounded off, protruding a flagellum, before
encysting; 9, young sporocyst; 10, the nucleus has divided into a brood of
eight; 11-14, stages of formation of zoospores. cv, Contractile vacuole; e,
mass of faecal granules; fl, flagellum; n, nucleus, × about 750⁄1.

Plasmodiophora infests the roots of Crucifers, causing the disease

known as "Hanburies," or "fingers and toes," in turnips, etc.
Serumsporidium dwells in the body cavity of small Crustacea. Many
of this group were described by Cienkowsky under the name of
"Monadineae" (in Arch. Mikr. Anat. i. 1865, p. 203). Zopf has added
more than anyone else since then to our knowledge. He
monographed them under Cienkowsky's name, as a subordinate
group of the Myxomycetes, "Pilzthiere oder Schleimpilze," in
Schenk's Handb. d. Bot. vol. iii. pt. ii. (1887). To Lankester (Encycl.
Brit., reprint 1891) we owe the name here adopted. Zopf has
successfully pursued their study in recent papers in his Beitr. Nied.
Org. The Chytridieae, usually ascribed to Fungi, are so closely allied
to this group that Zopf proposes to include at least the Synchytrieae
herein.

This group is very closely allied to Sporozoa; for the absence of

cytogamy, and of sickle-germs,[96] and of the complex spores and
cysts of the Neosporidia, are the only absolute distinctions.

6. Mycetozoa (Myxomycetes, Myxogastres)

Sarcodina moving and feeding by pseudopodia, with no skeleton,
aggregating more or less completely into complex "fructifications"
before forming 1-nucleate resting spores; these may in the first
instance liberate flagellate zoospores, which afterwards become
amoeboid, or may be amoeboid from the first; zoospores capable of
forming hypnocysts from which the contents escape in the original
form.

1. Aggregation taking place without

plastogamy, zoospores amoeboid, with a
clear ectosarc Acrasieae.
Copromyxa Zopf; Dictyostelium Brefeld.
2. Aggregation remaining lax, with merely
thread-like connexions, except when
encystment is to take place; cytoplasm
finely granular throughout; complete
fusion of the cytoplasm doubtful Filoplasmodieae
Labyrinthula Cienk.; Chlamydomyxa Archer; Leydenia (?)
Schaud.
3. Plasmodium formation complete, Myxomycetes.
 eventuating in the formation of a complex
fructification often traversed by elastic,
hygroscopic threads, which by their
contraction scatter the spores; zoospores
usually flagellate at first
Fuligo Hall.; Chondrioderma Rostaf.; Didymium Schrad. (Fig.
30).

I. The Acrasieae are a small group of saprophytes, often in the most

literal sense, though in some cases it has been proved that the
actual food is the bacteria of putrefaction. In them, since no cell-
division takes place in the fructification, it is certain that the
multiplication of the species must be due to the fissions of the
amoeboid zoospores, which often have the habit of Amoeba limax
(Fig. 1, p. 5).

II. Filoplasmodieae.—Chlamydomyxa[97] is a not uncommon

inhabitant of the cells of bog-mosses and bog-pools, and its nutrition
may be holophytic, as it contains chromoplasts; but it can also feed
amoeba-fashion. Labyrinthula is marine, and in its fructification each
of the component cells forms four spores. Leydenia has been found
in the fluid of ascitic dropsy, associated with malignant tumour.

III. Myxomycetes.—The fructification in this group is not formed by

the mere aggregation of the zoospores, but these fuse by their
cytoplasm to form a multinucleate body, the "plasmodium," which,
after moving and growing (with nuclear division) for some time like a
great multinucleate Reticularian, passes into rest, and develops a
fructification by the formation of a complex outer wall; within this the
contents, after multiplication of the nuclei, resolve themselves into
uninucleate spores, each with its own cyst-wall. The fructifications of
this group are often conspicuous, and resemble those of the
Gasteromycetous fungi (e.g., the Puffballs), whence they were at
first called Myxogastres. De Bary first discovered their true nature in
1859, and ever since they have been claimed by botanist and
zoologist alike.

The spore on germination liberates its contents as a minute

flagellate, with a single anterior lash and a contractile vacuole (Fig.
30, C). It soon loses the lash, becomes amoeboid, and feeds on
bacteria, etc. (Fig. 30, D, E). In this state it can pass into hypnocysts,
from which, as from the spores, it emerges as a flagellula. After a
time the amoeboids, which may multiply by fission, fuse on meeting,
so as to form the plasmodium (Fig. 30, F). This contains numerous
nuclei, which multiply as it grows, and numerous contractile
vacuoles. When it attains full size it becomes negatively hydrotactic,
crawls to a dry place, and resolves itself into the fructification. The
external wall, and sometimes a basal support to the fruit, are
differentiated from the outer layer of protoplasm; while the nuclei
within, after undergoing a final bipartition, concentrate each around
an independent portion of plasma, which again is surrounded as a
spore by a cyst-wall. Often the maturing plasmodium within the wall
of the fruit is traversed by a network of anastomosing tubes filled
with liquid, the walls of which become differentiated into membrane
like the fruit-wall, and are continuous therewith. As the fruit ripens
the liquid dries, and the tubes now form a network of hollow threads,
the "capillitium," often with external spiral ridges (Fig. 30, A, B).
These are very hygroscopic, and by their expansion and contraction
determine the rupture of the fruit-wall and the scattering of the
spores.