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 (above), boundary between pairing animals; (below), first fission; single
vertical line, continuity or enlargement. M, Meganucleus; µ, micronucleus;
Z, zygote-nucleus.

Fig. 54.—Four individuals of Coleps hirtus (Gymnostomaceae) swarming about

and ingesting a Vorticella (?) (From Verworn.)

The meganucleus lengthens, becomes irregularly constricted, and

breaks up into fragments, which are ultimately extruded or partially
digested. The micronucleus enlarges (Fig. 52, A) and undergoes
three successive divisions, or, strictly speaking, two fissions
producing four nuclei, of which one only undergoes the third. The
other three nuclei of the second fission degenerate like the
meganucleus.[167] Of the two micronuclei of this last division one
remains where it is as a "stationary" pairing nucleus, while its sister
passes as a "migratory" pairing-nucleus into the other mate, and
fuses with its stationary pairing-nucleus. Thus in either mate is
formed a "zygote-nucleus," or "fusion-nucleus." All these processes
are simultaneous in the two mates; and the migratory nuclei cross
one another on the bridge of junction of the two mates (Fig. 52, C).
Each mate now has its original cytoplasm (subject to the qualification
above), but its old nuclear apparatus is replaced by the fusion-
nucleus. This new nucleus undergoes repeated fissions; its offspring
enlarge unequally, the larger being differentiated as mega-, the
smaller as micro-nuclei. The mates now separate (Fig. 52, F, G), and
by the first (or subsequent) fission of each, the new mega- and
micro-nuclei are distributed to the offspring. Colpidium colpoda offers
the simplest case, on which we have founded our diagram showing
the nuclear relations. During conjugation the oral apparatus often
atrophies, and is regenerated; and in some cases the pellicle and
ciliary apparatus are also "made over."

Fig. 55.—Paramecium caudatum (Aspirotrichaceae). A, The living animal from

the ventral aspect; B, the same in optical section, the arrows show the
course taken by food-particles. buc.gr, Buccal groove; cort, cortex; cu,
cuticle; c.vac, contractile vacuole; f.vac, food vacuole; gul, gullet; med,
medulla; mth, mouth; nu, meganucleus; pa.nu, micronucleus; trch,
trichocysts discharged. (From Parker's Biology.)

In the Peritrichaceae the mates are unequal; the larger is the normal
cell, and is fixed; the smaller, mobile, is derived from an ordinary
individual by brood-divisions, which only occur under the conditions
that induce conjugation (Fig. 60). Here, though the two pairs of
nuclei are formed, it is only the migratory nuclei that unite, the
stationary ones aborting in both mates. During the final processes of
conjugation the smaller mate is absorbed into the body of the larger,
and so plays the part of male there. But this process, though one of
true binary sex, is clearly derived from the peculiar type of equal
reciprocal conjugation of the other Infusoria.
The Ciliata are almost all free-swimming animals with the exception
of most of the Peritrichaceae, and of the genera we now cite.
Folliculina forms a sessile tube open at either end; and Schizotricha
socialis inhabits the open mouths of a branching gelatinous tubular
stem, obviously secreted by the hinder end of the animal, and forking
at each fission to receive the produce. A similar habit to the latter
characterises Maryna socialis; all three species are marine, and
were described by Gruber.[168] Stentor habitually attaches itself by
processes recalling pseudopodia, and often forms a gelatinous
sheath.

The majority of the Oligotrichaceous Tintinnidae inhabit free

chitinous tests often beautifully fenestrated, as in Dictyocystis.

Many genera are parasitic in the alimentary canal of various

Metazoa, but none appear to be seriously harmful except
Ichthyophtheirius, which causes an epidemic in fresh-water fish.
Quite a peculiar fauna inhabit the paunch of Ruminants. Nyctotherus
and Balantidium are occasionally found in the alimentary canal of
Man.[169]

The Gymnostomaceae are predaceous, feeding for the most part on

smaller Ciliates. We have described the peculiar character of the
mouth and pharynx in this group, and the mail-like pellicle of Coleps
(Fig. 54). Loxophyllum is remarkable for the absence of cilia from
one of the sides of its flattened body, and the tufts of trichocysts
studding its dorsal edge at regular intervals. Actinobolus has
numerous tentacles, exsertile and retractile, each bearing a terminal
tuft of trichocysts, which serve to paralyse such active prey as
Halteria. Ileonema has one tentacle overhanging the mouth; and
Mesodinium has four short sucker-like projections around it.[170] It
has only two girdles of cilia, which are stout and resemble fine-
pointed cirrhi. In Dysteria the cilia are exclusively ventral, and the
naked dorsal surface has its pellicle condensed into a bivalve shell; a
posterior motile process ("foot") and a complex pharyngeal armature
add to the exceptional characters of the genus.

The Aspirotrichaceae are well known to every student of

"Elementary Biology" by the "type" Paramecium (Fig. 55), so
common in infusions, especially when containing a little animal
matter. P. bursaria often contains in its endosarc the green symbiotic
Flagellate Zoochlorella. Colpoda cucullus, very frequent in vegetable
infusions, usually only divides during encystment, and forms a brood
of four. Pleuronema chrysalis (Fig. 57) is remarkable for its habit of
lying for long periods on its side and for its immense undulating
membrane, forming a lip on the left of its mouth; Glaucoma has two,
right and left.

Fig. 56.—Trachelius ovum. A, general view; B, section through sucker; C, section

through contractile vacuole and its pore of discharge. al, Alveolar layer of
ectoplasm; cil, cilia; c.v, contractile vacuole; m, mouth; N, meganucleus; s,
sucker, from which pass inwards retractile myonemes. (After Clara
Hamburger.)
 Fig. 57.—Pleuronema chrysalis (Aspirotrichaceae). A, Unstimulated, lying quiet;
B, stimulated, in the act of springing by the stroke of its cilia. (From
Verworn.)

The Heterotrichaceae present very remarkable forms. Spirostomum

is nearly cylindrical, and, a very giant, may attain a length of 4 mm.
(1⁄6"). Stentor can attach itself by its hinder end, which is then finely
tapered and prolonged into a few pseudopodia; its body is trumpet-
shaped, with a spiral peristome forming a coil round its wide end,
and leading on the left side into the mouth. Many species when
attached secrete a gelatinous sheath or tube. S. polymorphus is
often coloured green by Zoochlorella (p. 125); S. coeruleus[171] and
S. igneus owe their names to the brilliant pigment, blue or scarlet,
deposited in granules in lines between the conspicuous longitudinal
myonemes. From their large size and elongated meganucleus
accompanied by numerous micronuclei, these two genera have
frequently been utilised for experiments on regeneration. In Metopus
sigmoides the peristomial area forms a dome above its wreath of
membranellae; and in M. pyriformis this is so great as to form the
larger part of the cell, which is top-shaped, tapering behind to a
point. Caenomorpha (Fig. 58) has the same general form, with a
peg-like tail, and possesses a girdle of cirrhi.[172] The converse
occurs in Bursaria; the cell is a half ellipse, something like a common
twin tobacco-pouch when closed: a deep depression thus occupies
the whole ventral surface, and opens by a wide slit extending along
the anterior end. The peristomial area occupies the dorsal side of the
pocket so formed, and the mouth is in the hinder left-hand corner.
Blepharisma sp. is parasitic in the Heliozoon Raphidiophrys viridis
(Fig. 20, 1, p. 74).
 Fig. 58.—Caenomorpha uniserialis. crh, Zone of cirrhi; c.t, cilia of tail; c.v,
contractile vacuole; c.w, ciliary wreath; g, granular aggregate; m, zone of
membranellae; N, meganucleus; n, micronucleus; oe, pharynx; t, tail-spine;
t1, accessory spine; u.m, undulating membrane; v, vacuole; z, precaudal
process. (After Levander.)

Among Oligotrichaceae, Halteria, common among the débris at the

bottom of pools in woods containing dead leaves, is remarkable for
an equatorial girdle of very long fine setae, and for its rapid erratic
darting movements, alternating with a graceful bird-like hover. The
Tintinnidae are mostly marine, pelagic, with the general look of a
stalkless Vorticella; some have a latticed chitinous shell.[173]

Fig. 59.—Stentor polymorphus. I, Young individual attached, extended; II, adult in

fission, contracted; cv in I, afferent canal of contractile vacuole; in II,
contractile vacuole; N, moniliform meganucleus (micronuclei omitted); o,
mouth; the fine lines are the myoneme fibrils. (From Verworn.)
Among Peritrichaceae, Vorticella (Fig. 60) and its allies have long
been known as Bell-animalcules to every student of pond-life. The
body has indeed the form of an inverted bell, closed at its mouth by
the "peristome," or oral disc; this is a short, inverted truncate cone
set obliquely so that its wide base hardly projects at one side, but is
tilted high on the other; the edge of the bell is turned out into a rim or
"collar," separated from the disc by a deep gutter. The collar,
habitually everted, or even turned down, contracts over the retracted
disc when the animal is retracted (E2), which is brought about by any
sort of shock, or when it swims freely backwards. For the latter
purpose a posterior ring of cilia (or rather membranellae) is
developed round the hinder end of the bell (A, cr, E3). The cilia of the
adoral wreath are very strong, united at the base into a continuous
membrane, and indeed themselves partake of the composite nature
of membranellae. The wreath forms more than one turn of a right-
handed spiral, the innermost turn ending abruptly on the disc, the
outer leading down into the mouth at the point where the disc is most
tilted and the groove deepest.[174] The pharynx (p) is long, and
contains an undulating membrane (u.m) on its inner side projecting
out through the mouth, and numerous cilia; it leads deep into the
body (p). The first part is distinguished as the "vestibule" (v), as into
it opens the anus, and the contractile vacuole (c.v.), the latter
sometimes opening by a reservoir (r). The body contains in the
ectoplasm myonema-fibrils which, by their contraction, withdraw the
disc, and at the same time circular fibrils close the peristome over it.
In the type-genus the pellicle is continued into a long, slender elastic
stalk (s), of which the longitudinal myoneme fibrils of the ectoplasm
converge to the stalk, and are prolonged into it as a spirally winding
fibre, sometimes transversely striated.[175] The effect of the
contraction of this is to pull the stalk into a helicoid spiral (like a coil-
spring), with the line of insertion of the muscle along the inner side of
the coils, which is, of course, the shortest path from one end to the
other (Fig. 60, B).
 Fig. 60.—Vorticella. A, expanded; B, stalk in contraction; c, eversible collar below
peristome; cr, line of posterior ciliary ring; c.v, contractile vacuole; m,
muscle of stalk; N, meganucleus; n, micronucleus; p, pharynx; r, reservoir of
contractile vacuole; s, tubular stalk; u.m, undulating membrane in vestibule;
v, hinder end of vestibule. E1, E2, two stages in binary fission; E3, free
zooid, with posterior wreath; F1, F2 division into mega- and micro-zooids
(m); G1, G2, conjugation; m, microzooid. (Modified from Bütschli, from
Parker and Haswell.)

The members of the Vorticellidae are very commonly attached to

weeds or to various aquatic Metazoa, each species being more or
less restricted in its haunts. Vorticella, the type, is singly attached to
a contractile stalk; fission takes place in the vertical plane, and one
of the two so formed retains the original stalk, while the other swims
off (Fig. 60, E1-E3), often to settle close by, so that the individuals
are found in large social aggregates, side by side, fringing water-
weeds with a halo visible to the naked eye, which disappears on
agitation by the sudden contraction of all the stalks. Carchesium and
Zoothamnium differ from Vorticella in the fact that the one daughter-
cell remains attached by a stalk coming off a little below the body of
the other, so as to give rise to large branching colonies.

In Carchesium (Fig. 51) the muscular threads of each cell are

separate, while in Zoothamnium they are continuous throughout the
colony. Epistylis has a solid, rigid stalk, and may give rise to
branching colonies, which often infest the body of the Water-Fleas
(Copepoda) of the genus Cyclops. Opercularia is characterised by
the depth of the gutter, the height of the collar, and the tapering
downward of the elongated disc. Vaginicola, Pyxicola, Cothurnia,
Scyphidia, all inhabit tubes, some of extreme elegance. Ophrydium
is a colonial form, found in ponds and ditches, resembling
Opercularia, but inhabiting tubes of jelly[176] that coalesce by their
outer walls into a large floating sphere; it usually contains the green
symbiotic Flagellate Zoochlorella. Trichodina is free, short, and
cylindrical, with both wreaths permanently exposed, and is provided
with a circlet of hooks within the aboral wreath. It is often parasitic, or
perhaps rather epizoic, on the surface of Hydra (see p. 254), gliding
over its body[177] with a graceful waltzing movement; it occurs also in
the bladder and genito-urinary passages of Newts, and even in their
body-cavity and kidneys.

II. Suctoria = Tentaculifera

Infusoria with cilia only in the young state,[178] without mouth or

anus, but absorbing food (usually living Ciliates) by one or more
tentacles, perforated at the apex; mostly attached, frequently epizoic,
rarely parasitic in the interior of other Protozoa.

Acineta, Ehrb. (Fig. 61, 2); Amoebophrya, Koppen; Choanophrya, Hartog (Fig.
62); Dendrocometes, St. (Fig. 61, 4); Dendrosoma, Ehrb. (Fig. 61, 9);
Endosphaera, Engelm.; Ephelota, Str. Wright (Fig. 61, 5, 8); Hypocoma,
Gruber; Ophryodendron, Cl. and L. (Fig. 61, 7); Podophrya, Ehrb. (Fig. 61, 1);
Rhyncheta, Zenker (Fig. 61, 3); Sphaerophrya, Cl. and L. (Fig. 61, 6),
Suctorella, Frenzel; Tokophrya, Bütschli.

This group, despite a superficial resemblance to the Heliozoa, show

a close affinity to the Ciliata; the nuclear apparatus is usually double
though a micronucleus is not always seen; the young are always
ciliated, and the mode of conjugation is identical in all cases hitherto
studied. Most of the genera are attached by a chitinous stalk (Fig.
61), continued in Acineta into a cup or "theca" surrounding the cell.
The pellicle is firm, often minutely shagreened or "milled" in optical
section by fine radial processes, whether superficial rods or the
expression of the meeting edges of radial alveoli is as yet uncertain.
The pellicle closely invests the ectosarc, is continued down into a
tubular sheath, from the base of which the tentacle rises, and
upwards to invest the tentacle, and is even prolonged into its cavity
in Choanophrya, the only genus where the tentacles are large
enough for satisfactory demonstration. These organs may be one or
more, and vary greatly in character. They may be (1) pointed for
prehension, puncture, and suction (Ephelota, Fig. 61, 5); (2) nearly
cylindrical, with a slightly "flared" truncate apex (Podophrya, Fig. 61,
1a); (3) filiform with a terminal knob; (4) "capitate" (Acineta, Fig. 61,
2); (5) bluntly truncate and capable of opening into a wide funnel for
the suction of food[179] (Choanophrya, Fig. 62; Rhyncheta, Fig. 61,
3). Their movements, too, are varied, including retraction and
protrusion, and a degree of flexion which reaches a maximum in
Rhyncheta (Fig. 61, 3), whose tentacle is as freely motile as an
elephant's trunk might be supposed to be were it as slender in
proportion to its length. They are continued into the body, and in
Choanophrya may extend right across it. In Podophrya trold the
pellicle rises into a conical tube about the base of the tentacle, which
is retracted through it completely with the prey in deglutition. In
Dendrocometes, Dendrosoma, and Ophryodendron (Fig. 61, 4, 9, 7),
the tentacles arise from outgrowths of the cell-body.
 Fig. 61.—Various forms of Suctoria. 1, a and b, two species of Podophrya; c, a
tentacle much enlarged; 2, a, Acineta jolyi; 2, b, A. tuberosa, with four
ciliated buds; in 6 the animal has captured several small Ciliata; 8, a, a
specimen multiplying by budding; 8, b, a free ciliated bud; 9, a, the entire
colony; 9, b, a portion of the stem; 9, c, a liberated bud. a, Organism
captured as food; b.c, brood-cavity; bd, bud; c.vac, contractile vacuole; l,
test; mg.nu, meganucleus; mi.nu, micronucleus; nu, nucleus; t, tentacle.
(From Parker and Haswell, after Bütschli and Saville Kent.)

The mechanism of suction is doubtful; but from the way particles

from a little distance flow into the open funnels of Choanophrya, it
may be the result of an increase of osmotic pressure. The external
pellicle of the tentacles is marked by a spiral constriction,[180] which
may be prolonged over the part included in the sheath. The
endosarc is rich in oil-drops, often coloured, and in proteid granules
which sometimes absorb stains so readily as to have been named
"tinctin bodies." It usually contains at least one contractile vacuole.

In Dendrocometes (and perhaps others) the whole cell may become

ciliated, detach itself and swim off; this it does when its host
(Gammarus) moults its cuticle.

In fission or budding we have to distinguish many modes. (1) In the

simplest, after the nuclear apparatus has divided, the cell divides
transversely; the distal half acquires cilia and swims off to attach
itself elsewhere, while the proximal remains attached. The tentacles
have previously disappeared and have to be formed afresh in both.
(2) More commonly fission passes into budding on the distal face; a
sort of groove deepens around a central prominence which becomes
the ciliated larva (Fig. 62, em); the tentacles of the "parent" are
retained. This process passes into (3) "internal budding," where a
minute pit leads into a bottle-shaped cavity.[181] (4) Again, the
budding may be multiple, the meganucleus protruding a branch for
each bud, while the micronucleus, by successive divisions, affords
the supply requisite. Sphaerophrya (Fig. 61, 6) and Endosphaera
multiply freely by fission within their Ciliate hosts, and were indeed
described by Stein as stages in their life-cycle. Conjugation of the
same type as in most Ciliates has been fully worked out in
Dendrocometes alone, by Hickson,[182] who has found the
meganuclei (though destined to disorganisation) conjugate for a
short time by the bridge of communication before the reciprocal
conjugation of the micronuclei.

We have referred to the endoparasitism of two genera.

Amoebophrya lives in several Acanthometrids, and in the aberrant
Radiolarian Sticholonche (see p. 86). The attached species are
some of them indifferent to their base; others are only found on
Algae, or again only epizoic on special Metazoan hosts, or even on
special parts of these. Thus Rhyncheta is only found on the couplers
of the thoracic limbs of Cyclops, and Choanophrya on the ventral
surface of its head and the adjoining appendages.
 Fig. 62.—Choanophrya infundibulifera. A, adult; B-D, tentacles in action in
various stages; E, tentacle at rest; F, young, just settled down, a, a, a,
Tentacles in various stages of activity; c, central cavity; c.v, contractile
vacuole; em, ciliated embryo showing contractile vacuole and nucleus; f,
spiral ridge; m, muscular wall of funnel; n, nucleus; tr, opening of funnel. (A-
D, F, modified after Zenker; E, original.)

We owe our knowledge of this group to the classical works of

Ehrenberg, Claparède and Lachmann, Stein, R. Hertwig, and
Bütschli. Plate has shed much light on Dendrocometes, and Hickson
has studied its conjugation. Ischikawa[183] has utilised modern
histological methods for the cytological study of Ephelota
bütschliana. René Sand has written a useful, but unequal, and not
always trustworthy monograph of the Order,[184] containing an
elaborate bibliography.
 PORIFERA (SPONGES)

BY

IGERNA B. J. SOLLAS, B.Sc. (Lond.)

Lecturer on Zoology at Newnham College, Cambridge.

CHAPTER VII

PORIFERA (SPONGES)[185]

INTRODUCTION—HISTORY—DESCRIPTION OF H A L I C H O N D R I A
P A N I C E A AS AN EXAMPLE OF BRITISH MARINE SPONGES AND OF
E P H Y D A T I A F L U V I A T I L I S FROM FRESH WATER—DEFINITION—
POSITION IN THE ANIMAL KINGDOM.

Sponges occupy, perhaps, a more isolated position than any other

animal phylum. They are not only the lowest group of multicellular
animals, but they are destitute of multicellular relatives. They are all
aquatic and—with the exclusion of a few genera found in fresh water
—marine, inhabiting all depths from between tide marks to the great
abysses of the ocean. They depend for their existence on a current
of water which is caused to circulate through their bodies by the
activity of certain flagellated cells. This current contains their food, it
is their means of respiration, and it carries away effete matters.
Consequently sponges cannot endure deprivation of oxygenated
water except for short periods, and only the hardiest inhabit regions
where the supply is intermittent, as between tide marks. This also
renders useless attempts to keep specimens in tanks, unless the
water is frequently renewed.
The outward appearance of sponges has an exceptionally wide
range, so that it is difficult to give a novice any very definite picture of
what he is to expect when searching for these animals. This diversity
is in part due to the absence of organs of sufficient size to determine
the shape of the whole or limit its variation, that is to say, the
separate organs are of an order of size inferior to that of the entire
body. The animals are fixed or lie loose on the sea bottom; there are
in no case organs of locomotion, and again no sense-organs, no
segregated organs of sex, and as a rule no distinction into axis and
lateral members. It is by these negative characters that the collector
may easily recognise a sponge.

History.—Sponges are, then, in many of their characters unique;

and they present a variety of problems for solution, both of special
and general interest, they are widely distributed in time and space,
and they include a host of forms. It therefore causes no little surprise
to learn that they have suffered from a long neglect, even their
animal nature having been but recently established. Though known
to naturalists from the time of Aristotle, sponges have been left for
modern workers as a heritage of virgin soil: it has yielded to them a
rich harvest, and is as yet far from exhausted.

The familiar bath sponge was naturally the earliest known member of
the phylum. It is dignified by mention in the Iliad and in the Odyssey,
and Homer, in his choice of the adjective "full of holes," πολύτρητος,
shows at least as much observation as many a naturalist of the
sixteenth and seventeenth centuries. Aristotle based his ideas of
sponges entirely upon the characters of the bath sponge and its near
allies, for these were the only kinds he knew. With his usual
perspicuity he reached the conclusion that sponges are animals,
though showing points of likeness to plants.

The accounts of sponges after Aristotle present little of scientific

interest until the last century. Doubtless this is in part due to the
absence of organs which would admit of dissection, and the
consequent necessity of finer methods of study. Like other attached
forms, sponges were plant or animal as it pleased the imagination of
the writer, and sometimes they were "plant animals" or Zoophyta:
those who thought them animal were frequently divided among
themselves as to whether they were "polypous" or "apolypous." An
opinion which it is somewhat difficult to classify was that of Dr.
Nehemiah Grew,[186] who says: "No Sponge hath any Lignous
Fibers, but is wholly composed of those which make the Pith and all
the pithy parts of a Plant, ... So that a Sponge, instead of being a
Zoophyton, is but the one-half of a Plant."

Sponges figure in herbals beside seaweeds and mushrooms, and

Gerarde says:[187] "There is found growing upon rockes near unto
the sea a certaine matter wrought together of the foame or froth of
the sea which we call Spunges ... whereof to speak at any length
would little benefit the reader ... seeing the use thereof is so well
known." About the middle of the eighteenth century, authors,
especially Peyssonnel, suggested that sponges were but the houses
of worms, which built them much as a bee or wasp builds nests and
cells. This was confuted by Ellis in 1765,[188] when he pointed out
that the sponge could not be a dead structure, as it gave proof of life
by "sucking and throwing out water." To Ellis, then, is due the credit
of first describing, though imperfectly, a current set up by sponges.
He mentions that Count Marsigli[189] had already made somewhat
similar observations.

It was not till 1825 that attention was again turned to the current,
when Robert Grant approached the group in a truly scientific
manner, and was ably supported by Lieberkühn. It would be
impossible to do justice to Grant in the brief summary to which we
must limit ourselves. The most important of his contributions was the
discovery that water enters the sponge by small apertures scattered
over the surface, and leaves it at certain larger holes, always
pursuing a fixed course. He made a few rough experiments to
estimate the approximate strength of the current, and, though he
failed to detect its cause, he supposed that it was probably due to
ciliary action. Grant's suggestion was afterwards substantiated by
Dujardin (1838), Carter (1847), Dobie (1852), and Lieberkühn
(1857). These five succeeded in establishing the claims of sponges
to a place in the animal kingdom, claims which were still further
confirmed when James-Clark[190] detected the presence of the
protoplasmic collar of the flagellated cells (see pp. 171, 176). Data
were now wanted on which to base an opinion as to the position of
sponges within the animal kingdom. In 1878 Schulze[191] furnished
valuable embryological facts, in a description agreeing with an earlier
one of Metschnikoff's, of the amphiblastula larva (p. 226) and its
metamorphosis. Then Bütschli[192] (1884) and Sollas[193] on
combined morphological and embryological evidence (1884)
concluded that sponges were remote from all the Metazoa, showing
bonds only with Choanoflagellate Protozoa (p. 121). This the exact
embryological work of Maas, Minchin, and Delage has done much to
prove, but it has to be admitted that unanimity on the exact position
of the phylum has not yet been attained, some authorities, such as
Haeckel, Schulze, and Maas still wishing to include sponges in the
Metazoa.

In this short history we have been obliged to refer only to work

helping directly to solve the problem of the nature of a sponge,
hence many names are absent which we should have wished to
mention.

Halichondria panicea.
One of the commonest of British sponges, which may be picked up
on almost any of our beaches, and which has also a cosmopolitan
distribution, is known by the clumsy popular name of the "crumb of
bread sponge," alluding to its consistency; or by the above technical
name, with which even more serious fault may be found.[194]

In its outward form H. panicea affords an excellent case of a

peculiarity common among sponges. Its appearance varies
according to the position in which it has lived. In fact, Bowerbank
remarks that it has no specific form. It may grow in sheets of varying
thickness closely attached to a rock, when it is "encrusting," or it is
frequently massive and lying free on the sea bottom; again, it may be
fistular, consisting of a single long tube, or it may be ridge-like,
apparently in this case consisting of a row of long tubes fused
laterally. In this last form it used to be called the "cockscomb
sponge," having been taken for a distinct species.

Bidder has proposed to call the different forms of the same species
"metamps" of the species. Figures of the metamps of H. panicea will
be found in Bowerbank's useful Monograph.[195]

The colour of the species is as inconstant as its form, ranging from

green to light brown and orange. MacMunn concludes from
spectroscopic work that H. panicea, contains at least three pigments,
a chlorophyll, a lipochrome, and a histohaematin.[196] Lipochromes
vary from red to yellow, chlorophyll is always associated with one or
more of them. Histohaematin is a respiratory pigment. Proof has not
yet been adduced that the chlorophyll is proper to the sponge and is
not contained in symbiotic algae.

In spite of all this inconstancy H. panicea, is one of the most easily

determined species. It is only necessary to dry a small fragment,
including the upper surface; a beautiful honeycomb-like structure is
then visible on this surface, and among British sponges this is a
property peculiar to the species (Bowerbank). Whatever the form of
the sponge, one or more large rounded apertures are always present
on the exterior; these are the "oscula." In the encrusting metamp the
oscula are flush with the general surface, while in the other cases
they are raised on conical projections; fistular specimens carry the
osculum at the distal end, and the cockscomb has a row of them
along its upper edge. Much more numerous than the oscula are
smaller apertures scattered over the general surface of the sponge,
and known as "ostia."
 Fig. 63.—Portion of the surface of H. panicea, from dried specimen. A, natural
size; B, magnified. The large shaded patches are ostia.

If the sponge be placed in a shallow glass dish of sea water the

function of the orifices can be made out with the naked eye,
especially if a little powdered chalk or carmine be added to the water.
If the specimen has been gathered after the retreating tide has left it
exposed for some time, this addition is unnecessary, for as soon as it
is plunged into water its current bursts vigorously forth, and is
rendered visible by the particles of detritus that have accumulated in
the interior during the period of exposure and consequent
suspended activity. The oscula then serve for the exit of currents of
water carrying particles of solid matter, while the entrance of water is
effected through the ostia.

Sections show that the ostia lead into spaces below the thin
superficial layer or "dermal membrane"; these are continued down
into the deeper parts of the sponge as the "incurrent canals,"
irregular winding passages of lumen continually diminishing as they
descend. They all sooner or later open by numerous small pores
—"prosopyles"—into certain subspherical sacs termed flagellated
chambers. Each chamber discharges by one wide aperture
—"apopyle"—into an "excurrent canal." This latter is only
distinguishable from an incurrent canal by the difference in its mode
of communication with the chambers.
 Fig. 64.—H. panicea: the arrows indicate the direction of the current, which is
made visible by coloured particles. (After Grant.)

The excurrent canals convey to the osculum the water which has
passed through the ostia and chambers. All the peripheral parts of
the sponge from which chambers are absent are termed the
"ectosome," while the chamber-bearing regions are the
"choanosome."

The peculiar crumb-of-bread consistency is due to the nature of the

skeleton, which is formed of irregular bundles and strands of minute
needles or spicules composed of silica hydrate, a substance familiar
to us in another form as opal: they are clear and transparent like
glass. They are scattered through the tissues in great abundance.

The classes of cellular elements in the sponge are as follows:

Flattened cells termed "pinacocytes" cover all the free surfaces, that
is to say, the external surface and the walls of the excurrent and
incurrent canals. The flagellated chambers are lined by
"choanocytes" (cf. Fig. 70, p. 176); these are cells provided at their
inner end with a flagellum and a collar surrounding it. They resemble
individuals of the Protozoan sub-class Choanoflagellata, and the
likeness is the more remarkable because no other organisms are
known to possess such cells. Taken together the choanocytes
constitute the "gastral layer," and they are the active elements in
producing the current. The tissue surrounding the chambers thus
lying between the excurrent and incurrent canals consists of a
gelatinous matrix colonised by cells drawn from two distinct sources.
In the first place, it contains cells which have a common origin with
the pinacocytes, and which together with them make up the "dermal