Protozoology 1946 Kudo
Protozoology 1946 Kudo
Protozoology 1946 Kudo
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PROTOZOOTvOGY
(^rd Edition)
PROTOZOOLOGY
By
Professor of Zoology
The University of Illinois
Urbana, Illinois
CHARLES C THOMAS •
PUBLISHER
EAST LAWRENCE AVENUE
301-327
SPRINGFIELD ILLINOIS
•
1946
Published by Charles C Thomas
301-327 East Lawrence Avenue, Springfield, Illinois
now given.
Chapters 1 to 6 deal with introduction, ecology, morphology,
physiology, reproduction, and variation and heredity, of Proto-
zoa. Each subject-matter has been considered in the light of more
recent investigations as fully as the space permitted. Selection of
material from so great a number of references has been a very diffi-
cult task. If any important papers have been omitted, it was en-
tirely through over-sight on the part of the author.
The taxonomic portion (Chapters 7 to 43) has also been com-
pletely rewritten and enlarged. Numerous genera and species, both
old and new, have been added synonymy of genera and species has
;
1935) however, unlike the latter, all parasitic ciliates have also been
;
59941
CONTENTS
9 Order 2 Cryptomonadina 213
10 Order 3 Phytomonadina 217
11 Order 4 Euglenoidina 232
Order 5 Chloromonadina 243
12 Order 6 Dinoflagellata 245
13 Subclass 2 Zoomastigina 263
Order 1 Rhizomastigina 263
14 Order 2 Protomonadina 268
15 Order 3 Polymastigina 293
16 Order 4 Hypermastigina 318
17 Class 2 Sarcodina 328
Subclass 1 Rhizopoda 329
Order 1 Proteomyxa 329
18 Order 2 Mycetozoa 335
19 Order 3 Amoebina 343
20 Order 4 Testacea 374
21 Order 5 Foraminifera 394
22 Subclass 2 Actinopoda 406
Order 1 Heliozoa 406
23 Order 2 Radiolaria 417
24 Class 3 Sporozoa 427
Subclass 1 Telosporidia 427
Order 1 Gregarinida 428
25 Order 2 Coccidia 464
26 Order 3 Haemosporidia 484
27 Subclass 2 Acnidosporidia 507
Order 1 Sarcosporidia 507
Order 2 Haplosporidia 510
28 Subclass 3 Cnidosporidia 515"
Introduction
sidered that the binary fission of the Protozoa and of the Protophyta
is and transverse, respectively. A great majority of
longitudinal
Ciliata, however, multiply by transverse division. In general the
nutrition of Protozoa is holozoic and of Protophyta, holophytic;
but there are large numbers of Protozoa which nourish themselves
by holophytic method. Thus an absolute and clean-cut separation
of the two groups of unicellular organisms is not possible. Haeckel
6 PROTOZOOLOGY
coined the name
Protista to include these organisms in a single
group, but this not generally adopted, since it includes undoubted
is
both the hosts and the parasites, than to assume that there once
existed between Patagonia and Australia a land connection over
which frogs, containing Zelleriella, migrated.
Experimental studies of large Protozoa have thrown light on the
relation between the nucleus and the cytoplasm, and have furnished
a basis for an understanding of regeneration in animals. In Protozoa
we find various types of nuclear divisions ranging from a simple
amitotic division to a complex process comparable in every detail
with the typical metazoan mitosis. A part of our knowledge in
cytology is based upon studies of Protozoa.
Through the efforts of various investigators in the past fifty
years, it has now become known
that numerous parasitic Protozoa
occur in man
(Kudo, 1944). Entamoeba histolytica, Balantidium coli,
and three species of Plasmodium, all of which are pathogenic to man,
are widely distributed throughout the world. In certain restricted
areas are found other pathogenic forms, such as Trypanosoma and
Leishmania. Since all parasitic Protozoa presumably have originated
in free-living forms and since our knowledge of the morphology,
physiology, and reproduction of the parasitic forms has largely been
obtained in conjunction with the studies of the free-living organ-
knowledge of the entire phylum is necessary to under-
isms, a general
stand the parasitic forms.
Recent studies have further revealed that almost all domestic
animals are hosts to numerous parasitic Protozoa, many of which
INTRODUCTION 9
occur. On the other hand, other Microsporidia are now known to in-
fect certain insects, such as mosquitoes and lepidopterous pests,
which, when heavily infected, die sooner or later. Methods of de-
struction of these insects by means of chemicals are more and more
used, but attention should also be given to utilization of the parasitic
Protozoa and Protophyta for this purpose.
While the majority of Protozoa lack permanent skeletal structures
and their fossil forms are unknown, there are at least two large
groups in the Sarcodina which possess conspicuous shells and which
are found as fossils. They are Foraminifera and Radiolaria. From
early palaeozoic era down to the present day, the carbonate of
lime which makes up the skeletons of numerous Foraminifera has
been left embedded in various rock strata. Although there is no dis-
tinctive foraminiferan fauna characteristic of a given geologic pe-
riod, there are certain peculiarities of fossil Foraminifera which dis-
tinguish one formation from the other. From this fact one can un-
derstand that knowledge of foraminiferous rocks is highly useful in
checking up logs in well drilling. The skeletons of the Radiolaria are
the main constituent of the ooze of littoral and deep-sea regions.
They have been found abundantly in siliceous rocks of the palaeozoic
and the mesozoic eras, and are also identified with the clays and
other formations of the miocene period. Thus knowledge of these two
orders of Sarcodina, at least, is essential for the student of geology
and paleontology.
and interesting forms. This work was followed by Stokes' The Fresh-
water Infusoria of the United States, which appeared in 1888.
Butschli (1880-1889) established Sarcodina and made an excellent
contribution to the taxonomy of the then-known species of Protozoa,
which is still considered as one of the most important works on gen-
eral protozoology. The painstaking researches by Maupas, on the
conjugation of ciliates, corrected erroneous interpretation of the
phenomenon observed by Balbiani some thirty years before and gave
impetus to a renewed cytological study of Protozoa. The variety in
form and structure of the protozoan nuclei became the subject of in-
tensive studies by several cytologists. Weismann (1881) put into
words the immortality of the Protozoa. Schaudinn contributed much
toward the cytological and developmental studies of Protozoa.
In the first year of the present century, Calkins in the United
States and Doflein in Germany wrote modern textbooks on protozo-
ology dealing with the biology as well as the taxonomy. Calkins
initiated the so-called isolation pedigree culture of ciliates in order to
study the physiology of conjugation and other phenomena connected
with the life-history of the ciliates. Recently application of bacteria-
free culture technique to certain free-living flagellates and ciliates has
brought to light hitherto unknown facts regarding nutritional re-
quirements of these organisms.
Today the Protozoa are more and more intensively and exten-
sively studied from both the biological and the parasitological sides,
and important contributions appear continuously. Since all parasitic
Protozoa appear to have originated in free-living forms, the com-
prehension of the morphology, physiology, and development of the
latter group is obviously fundamentally important for a thorough
understanding of the former group.
Compared with the advancement of our knowledge on free-living
Protozoa, that on parasitic forms has been very slow. This is to be ex-
pected, of course, since the vast majority of them are so minute that
the discovery of their presence has been made possible only through
improvements in the microscope and in technique.
Here again Leeuwenhoek seems to have been the first to observe
a parasitic protozoan, for he observed, according to Dobell, in the
fall of 1674, the oocysts of the coccidian, Eimeria stiedae, in the con-
References
BtJTSCHLi, O. 1887-1889 Bronn's Klassen und Ordnungen des
Thier-reichs. Vol. 1, Part 3.
Calkins, G. N. 1933 The biology of the Protozoa. 2 ed. Philadelphia.
Cole, F. J. 1926 The history of protozoology London.
.
Ecology
17
18 PROTOZOOLOGY
and 7.6, excess of potassium decreases resistance and excess of cal-
cium increases resistance. Glaser and Coria (1933) cultivated Para-
mecium caudatum on dead yeast free from living organisms at
20-28°C. (optimum 25°C.) and noted that at 30°C. the organisms
were killed. Doudoroff (1936), on the other hand, found that in
P. multimicronucleatum its resistance to raised temperature was low
in the presence of food, but rose to a maximum when the food was
exhausted, and there was no appreciable difference in the resistance
between single and conjugating individuals.
The thermal waters of hot springs have been known to contain liv-
ing organisms including Protozoa. Glaser and Coria obtained from
the thermal springs of Virginia, several species of Mastigophora,
and an amoeba which were living in the water, the tempera-
Ciliata,
ture of whichwas 34-36°C., but did not notice any protozoan in the
water which showed 39-41°C. Uyemura and his co-workers made a
series of studies on Protozoa living in various thermal waters of Ja-
pan, and reported that many species lived at unexpectedly high
temperatures. Some of the Protozoa observed and the temperatures
of the water in which they were found are as follows: Amoeba sp.,
Vahlkampfia Umax, A. radiosa, 30-51°C.; Amoeba verrucosa, Chilo-
donella sp., Lionotus fasciola, Paraynecium caudatum, 36-40°C.;
Oxytricha fallax, 30-56°C.
Under experimental conditions, it has been shown repeatedly that
many protozoans become accustomed to a very high temperature if
the change be made gradually. Dallinger (1887) showed a long time
ago that Tetramitus rostratus and two other species of flagellates
became gradually acclimatized up to 70°C. in several years. In na-
ture, however, the thermal death point of most of the free-living
Protozoa appears to lie between 36° and 40°C. and the optimum
temperature, between 16° and 25°C.
On the other hand, the low temperature seems to be less detri-
mental to Protozoa than the higher one. Many protozoans have
been found to live in water under ice, and several haematochrome-
bearing Phytomastigina undergo vigorous multiplication on snow in
high altitudes, producing the so-called "red snow." Klebs (1893) sub-
jected the trophozoites of Euglena to repeated freezing without ap-
parent injury and Jahn (1933) found no harmful effect when Euglena
cultures were kept without freezing at — 0.2°C. for one hour, but
when kept at — 4°C. for one hour the majority were killed. Gaylord
(1908) exposed Trypanosoma gambiense to liquid air for 20 minutes
without apparent injury, but the organisms were killed after 40 min-
utes' immersion.
ECOLOGY 19
canal of the animal unharmed or are introduced after the faeces are
voided, and undergo development and multiplication in the faecal
infusion. Such forms are collectively called coprozoic Protozoa. The
coprozoic protozoans grow easily in suspension of old faecal matter
which are rich in decomposed organic matter and thus show a strik-
ingly strong capacity of adapting themselves to conditions different
from those of the water in which they normally live. Some of the
Protozoa which have been referred to as coprozoic and which are
mentioned in the present work are, as follows: Scytomonas pusilla,
Rhynchomonas nasuta, Cercomonas longicauda, C. crassicauda, Tre-
yomonas agilis, Dimastig amoeba gruheri, Acanthamoeba hyalina,
Chlamydophrys stercorea and Tillina magna.
As a rule, the presence of sodium chloride in the sea water prevents
the occurrence of the large number of fresh-water inhabitants. Cer-
tain species, however, have been known to live in both fresh and
brackish or salt water. Among the species mentioned in the present
work, the following species have been reported to occur in both fresh
and salt waters: Mastigophora: Amphidinium lacustris, Cerat-
ium hirundinella; Sarcodina: Lieberkuhnia wagneri; Ciliata: Meso-
dinium pulex, Prorodon discolor, Lacrymaria olor, Amphileptus
claparedei, Lionotus fasciola, Nassula aurea, Trochilioides recta,
Chilodonella cucullulus, Trimyema compressum, Paramecium cal-
kinsi, Colpidium campylum, Platynematum sociale, Cinetochilum
margaritaceum, Pleuronema coronatum, Caenomorpha medusula,
Spirostomum minus, S. teres, Climacostomum virens, and Thuricola
follicidata; Snctoria: Metacineta mystacina, Endosphaera engelmanni.
It seems probable that many other protozoans are able to live
in both fresh and salt water, judging from the observations such
as that made by Finley (1930) who subjected some fifty species of
freshwater Protozoa of Wisconsin to various concentrations of sea
water, either by by gradual addition of the sea
direct transfer or
water. He found that Bodo uncinatus, Uronema marina, Pleuron-
ema jaculans and Colpoda aspera are able to live and reproduce
even when directly transferred to sea water, that Amoeba verrucosa,
Euglena, Phacus, Monas, Cyclidium, Euplotes, Lionotus, Para-
mecium, Styl onychia, etc., tolerate only a low salinity when directly
transferred, but, if the salinity is gradually increased, they live in
100 per cent sea water, and that Arcella, Cyphoderia, Aspidisca, Ble-
pharisma, Colpoda cucullus, Halteria, etc. could not tolerate 10 per
cent sea water even when the change was gradual. Finley noted no
morphological changes in the experimental protozoans which might
be attributed to the presence of the salt in the water, except Amoeba
22 PROTOZOOLOGY
verrucosa, inwhich certain structural and physiological changes were
observed as follows: as the salinity increased, the pulsation of the
contractile vacuole became slower. The body activity continued up
to 44 per cent sea water and the vacuole pulsated only once in 40
minutes, and after systole, it did not reappear for 10-15 minutes.
The organism became less active above this concentration and in
84 per cent sea water the vacuole disappeared, but there was still a
tendency to form the characteristic ridges, even in 91 per cent sea
water, in which the organism was less fan-shaped and the cytoplasm
seemed to be more viscous. Yocom (1934) found that Eiiplotes pa-
tella was able to live normally and multiply up to 66 per cent of
ECOLOGY 23
pfiKppillTf
dren show usually a low and a high rate of malaria infection respect-
ively,but the latter frequently do not show symptoms of infection,
even though the parasites are detectable in the blood. Apparently
repeated infection produces tolerance which can keep, as long as the
host remains healthy, the parasites under control. There seems to be
immunity against Plasmodium
also racial difference in the degree of
and Trypanosoma, as shown by James, Milam and Kusch, etc.
As to the mechanism of immunity, the destruction of the parasites
by phagocytosis of the endothelial cells of the spleen, bone marrow
and liver and continued regenerative process to replace the de-
stroyed blood cells, are the two important phases in the cellular de-
fense mechanism. Besides, there are indications that humoral de-
fense mechanism through the production of antibodies is in active
operation in infections by Plasmodium knowlesi (Coggeshall and
Kumm, Eaton, etc.) and trypanosomes (Taliaferro),
With regard to the origin of parasitic Protozoa, it is generally
agreed among biologists that the parasite in general evolved from
the free-living form. The protozoan association with other organ-
isms was begun when various protozoans which lived attached to,
or by crawling submerged objects happened to transfer them-
on,
selves to various invertebrateswhich occur in the same water.
These Protozoa benefit by change in location as the host animal
moves about, and thus enlarging the opportunity to obtain a con-
tinued supply of food material. Such ectocommensals are found
abundantly; for example, the peritrichous ciliates attached to the
body and appendages of various aquatic animals such as larval in-
sects and microcrustaceans. Ectocommensalism may next lead to
ectoparasitism as in the case of Costia or Hydramoeba, and then
again instead of confining themselves to the body surface, the Pro-
tozoa may bore into the body wall from outside and actually acquire
the habit of feeding on tissue cells of the attached animals as in the
case of Ichthyophthirius.
The next step in the evolution of parasitism must have been
reached when Protozoa, accidentally or passively, were taken into
the digestive system of the Metazoa. Such a sudden change in
habitat appears to be fatal to most protozoans. But certain others
possess extraordinary capacity to adapt themselves to an entirely
different environment. For example, Dobell (1918) observed in the
tadpole gut, a typical free-living limax amoeba, with characteristic
nucleus, contractile vacuoles, etc., which was found in numbers in
the water containing the faecal matter of the tadpole. Glaucoma
pyriformis (Tetrahymena geleii), a free-living ciliate, was found to
30 PROTOZOOLOGY
occur in the body cavity of the larvae of Theohaldia annulata (after
MacArthur) and in the larvae of Chironomus -plumosus (after Treil-
lard and Lwoff). Lwoff successfully inoculated this ciliate into the
larvae of Galleria mellonella which died later from the infection.
Recently Janda and Jirovec (1937) injected bacteria-free culture of
intestine and other viscera of the land slug, Agriolimax agrestis, the
slug forms being much larger than the free-living individuals.
It may be further speculated that Vahlkampfia, Hydramoeba,
Schizamoeba, and Endamoeba, are the different stages of the course
the intestinal amoebae might have taken during their evolution.
Obviously endocommensalism in the alimentary canal was the
initial phase of endoparasitism. When these endocommensals began
to consume an excessive amount of food or to feed on the tissue cells
of the host gut, they became the true endoparasites. Destroying or
penetrating through the intestinal wall, they became first established
in the body or organ cavities and then invaded tissues, cells or even
nuclei, thus developing into pathogenic Protozoa. The endoparasites
ECOLOGY 31
References
Calkins, G. N. and F. M. Summers (editors). 1941 Protozoa in
biological research. New York.
Chalkley, H. W. 1930 Resistance of Paramecium to heat as af-
fected by changes in hydrogen-ion concentration and in inor-
ganic salt balance in surrounding medium. U. S. Publ. Health.
Rep., Vol. 45.
Cleveland, L. R. 1926 Symbiosis among animals with special
reference to termites and their intestinal flagellates. Quart. Rev.
Biol., Vol. 1.
1928 Tritrichomonas fecalis nov. sp. of man; its ability to
grow and multiply indefinitely in faeces diluted with tap water
and in frogs and tadpoles. Amer. Jour. Hyg., Vol. 8.
Dallinger, W. H. 1887 The president's address. Jour. Roy. Micr.
Soc. for 1887.
DoBELL, C. 1918 Are Entamoeba histolytica and Entamoeba ranarum
the same species? Parasitology, Vol. 10.
DouDOROFF, M. 1936 Studies in thermal death in Paramecium.
Jour. Exp. Zool., Vol. 72.
Efimoff, W. W. 1924 Ueber Ausfrieren und Ueberkaltung der
Protozoen. Arch. f. Protistenk., Vol. 49.
Finley, H. E. 1930 Toleration of freshwater Protozoa to increased
salinity. Ecology, Vol. 11.
Glaser, R. W. and ISF. A. Coria. 1933 The culture of Paramecium
caudatum free from living microorganisms. Jour. Parasit. Vol.
20.
Janda, V. and 0. Jirovec 1937 Ueber kiinstlich hervorgerufenen
Parasitismus eines f reilebenden Ciliaten Glaucoma piriformis und
Infektionsversuche mit Euglena gracilis und Spirochaeta hiflexa.
Mem. soc. zool tehee, de Prague, Vol. 5.
Kidder, G. W. 1941 Growth studies on cihates. VII. Biol. Bull.,
Vol. 80.
Kolkwitz, R. and M. Marsson 1909 Oekologie der tierischen
Saprobien. Intern. Rev. Ges. Hydrobiol. u. Hydrogr., Vol. 2.
Kudo, R. R. 1929 Histozoic Myxosporidia found in freshwater
fishes of Illinois, U.S.A. Arch. f. Protistenk., Vol. 65.
Lackey, J. B. 1925 The fauna of Imhof tanks. Bull. New Jersey
Agr. Exp. Stat., No. 417.
Lauterborn, R. 1901 Die "sapropelische" Lebewelt. Zool. Anz.,
Vol. 24.
Needhum, J. G., P. S. Galtsoff, F. E. Lutz and P. S. Welch. 1937
Culture methods for invertebrate animals. Ithaca, N.Y.
Noland, L. E. 1925 Factors influencing the distribution of fresh-
water ciliates. Ecology, Vol. 6.
32 PROTOZOOLOGY
Reynolds, B. D. 1936 Colpoda steini, a facultative parasite of the
land slug, Agriolimax agrestis. Jour. Parasit., Vol. 22.
and J. B. Looper 1928 Infection experiments with Hydra-
moeha hydroxena nov. gen. Ibid., Vol. 15.
Sandon, H. 1927 The composition and distribution of the protozoan
fauna of the soil. Edinburgh.
Taliaferro, W. H. 1926 Host resistance and types of infections in
trypanosomiasis and malaria. Quart. Rev. Biol., Vol. 1.
VON Brand, T. 1938 The metabolism of pathogenic trypanosomes
and the carbohydrate metabolism of their hosts. Ibid., Vol. 13.
Wenyon, C. M. 1926 Protozoology. 2 vols. London and New York.
WoLFSON, C. 1935 Observations on Paramecium during exposure
to sub-zero temperatures. Ecology, Vol. 16.
YocoM, H. B. 1934 Observations on the experimental adaptation
of certain freshwater ciliates to sea water. Biol. Bull., Vol. 67.
Chapter 3
Morphology
33
34 PROTOZOOLOGY
phase of the metabolism. Aside from the importance as the control-
ling center of metabolism, evidences point to the conclusion that the
nucleus contains the genes or hereditary factors which characterize
each species of protozoans from generation to generation, as in the
cells of multicellular animals and plants.
The nucleus
Because of a great variety of the body form and organization, the
protozoan nuclei are of various forms, sizes and structures. At one
extreme there is a small nucleus and, at the other, a large voluminous
one and, between these extremes, is found almost every conceivable
variety of form and structure. The majority of Protozoa contain a
single nucleus, though many may possess two or more throughout
the greater part of their life-cycle. In several species, each individual
possesses two similar nuclei, as in Diplomonadina, Protoopalina
and Zelleriella. In Euciliata and Suctoria, two dissimilar nuclei, a
macronucleus and a micronucleus, are typically present. The macro-
nucleus is always larger than the micronucleus, and controls the
trophic activities of the organism, while the micronucleus is con-
cerned with the reproductive activity. Certain Protozoa possess
numerous nuclei of similar structure, as for example, in Pelomyxa,
Mycetozoa, Actinosphaerium, Opalina, Cepedea, Myxosporidia,
Microsporidia, etc.
The essential components of the protozoan nucleus are the nuclear
membrane, chromatin, plastin and nucleoplasm. Their interrela-
tionship varies sometimes from one developmental stage to an-
other, and vastly among different species. Structurally, they fall in
general into one of the two types: vesicular and compact.
The vesicular nucleus (Fig. 2, a) consists of a nuclear membrane
which is sometimes very delicate but distinct, nucleoplasm and
chromatin. Besides there is an intranuclear body which is, as a rule,
more or less spherical and which appears to be of different make-ups
as judged by its staining reactions among different nuclei. It may be
composed of chromatin, of plastin, or of a mixture of both. The first
type is sometimes called karyosome and the second, nucleolus or
plasmosome. Absolute distinction between these two terms cannot
be made as they are based upon the difference in affinity to nuclear
stains which cannot be standardized and hence do not give uni-
formly the same result. Following Minchm and others, the term
endosome is advocated here to designate one or more conspicuous
bodies other than the chromatin granules, present within the nuclear
membrane.
MORPHOLOGY 35
Nuclear membrane
Endosome
Achromatic strand
Chromatin granules
a b
nuclei, since there are numerous nuclei the structures of which are
intermediate between the two. Moreover what appears to be a
vesicular nucleus in life, may approach a compact nucleus when
fixed and stained as in the case of Euglenoidina. Several experimental
observations show that the number, size, and structure of the endo-
MORPHOLOGY 37
The cytosome
The extranuclear part of the protozoan body is the cytosome.
It composed of the cytoplasm, a colloidal system, which may be
is
of plasticity.
The pellicle of a ciliate is much thickerand more definite, and
often variously ridged or sculptured. In many, linear furrows and
ridges run longitudinally, obliquely, or spirally; and, in others, the
ridges are combined with hexagonal or rectangular depressed areas.
Still in others,such as Coleps, elevated platelets are arranged paral-
lel to the longitudinal axis of the bodJ^ In certain peritrichous
ciliates, such as Vorticella monilata, Carchesiiim granulatum, etc.,
the pellicle may possess nodular thickenings arranged in more or less
parallel rows at right angles to the body axis.
While the pellicle always covers the protozoan body closely,
there are other kinds of protective envelopes produced by Protozoa
which may cover the body rather loosely. These are the shell, test,
lorica or envelope. The shell of various Phytomastigina is usually
made up of cellulose, a carbohydrate, which is widely distributed
40 PROTOZOOLOGY
in the plant kingdom. It may be composed of a single or several
layers, and may possess ridges or markings of various patterns on it.
nucleate forms, or from the zone between the ectoplasm and endo-
plasm (Fig. 6). Although semipermanent in structure, the axial rod
easily absorbed and reformed. In the genera of Heliozoa, not
is
' '7'
r-
5^®ID
dition of KOH
solution to the water
Anterior flagellum
Flagellum
Basal granule
Blepharoplast
Rhizoplast
Undulating Nucleus
membrane Parabasal body
Nucleus
Basal granule
posterior end of the body and would push the body forward by its
vibration. Lankester coined the terms tractella and pulsella for
pulling and pushing flagella respectively.
In certain parasitic Mastigophora, such as Trypanosoma (Fig.
9, a). Trichomonas, etc., there is a very delicate membrane extending
out from the side of the body, a flagellum bordering its outer margin.
When this membrane vibrates, shows a characteristic undulating
it
form much resembling the lipoid structure of Gelei and called them
"cross-striation" of the contractile component (Fig. 10).
MORPHOLOGY 49
Cirrus fiber
Ectoplasmic granules
Basal granules of
component cilia
Undulating membrane
cpg
pletely (Figs. 11, 12), which was demonstrated by Taylor. Klein also
showed by desiccation that each marginal cirrus of Stylonychia was
composed of 7 to 8 cilia. In some instances, the distal portion of a
cirrus may show two or more branches. The cirri are confined to the
ventral surface in Hypotricha, and called frontal, ventral, anal,
caudal, and marginal cirri, according to their location (Fig. 11). Un-
like the cilia, the cirri may move in any direction so that the organ-
isms bearing them, show various types of locomotion. Oxytricha,
MORPHOLOGY 51
n
i^^
jHOl'
Fibrillar structures
mc
1 ^^- bg
i*^:*/
MORPHOLOGY 53
oe
"
0^^<
p, pelhcle; prs, pharyngeal retractor strands; si, skeletal laminae; vs, ven-
tral skeletal area.
56 PROTOZOOLOGY
ectoplasm of the operculum (opercular fibers). A similar apparatus
has since been observed in many other ciliates: Euplotes CYocom;
Taylor), Balantiduum (McDonald), Paramecium (Rees; Brown;
Lund), Tintinnopsis Boveria (Pickard), Dileptus
(Campbell),
(Visscher), Chlamydodon (MacDougall), Entorhipidium and Le-
chriopyla (Lynch), Eupoterion (MacLennan and Connell), Metopus
(Lucas), Troglodytella (Robertson), Oxytricha (Lund), Ancistruma
and Conchophthirus (Kidder), etc.
along the proximal border of the oral lip and the bases of all mem-
branellae. Yocom further noticed that within the lip there is a
latticework structure whose bases very closely approximate the cyto-
stomal Taylor (1920) recognized two additional groups of
fiber.
which marks the outer end of the trichocyst. The second part which
Klein called "directly connected" (subpellicular) conductile system
consists essentially of the longitudinal lines connecting all basal
granules in a longitudinal row of hexagons and of delicate transverse
fibrils connecting granules of adjacent rows especially in the cyto-
J)
thr
•i ;:
W%s^
trb
t >
extr
trg
trb
rt
Hold-fast organellae
long spirally coiled delicate thread, the polar filament (Fig. 259).
66 PROTOZOOLOGY
The polar filament is considered as a temporary anchoring organella
of the spore at the time of its germination after it gained entrance
into the alimentary canal of a suitable host. In the Microsporidia,
the filament may or may not be enclosed within a capsule. The ne-
matocysts (Fig. 110, h) of certain dinoflagellates belonging to Nema-
toidium and Polykrikos, are almost identical in structure with those
The blepharoplast
In the Mastigophora or in other groups in which flagellate stages
occur, the flagellum ends internally in a basal granule, which, in
turn, is sometimes connected by a much larger bod}^ This latter
organella has been called the blepharoplast. In many instances
they appear to be combined in one. The blepharoplast is further
connected by a fibril, the rhizoplast, with the nucleus (Fig. 24).
The blepharoplast and centriole are considered synonymous by
Minchin, Cleveland, and others, since they give rise to the kinetic
organella.Woodcock and Minchin held, on the other hand, that the
blepharoplast was a nucleus holding a special relation with loco-
motor organellae, and called it kinetonucleus. In recent years it
has become known that the blepharoplast of many flagellates re-
68 PROTOZOOLOGY
spends positively to Feulgen's nucleal reaction which may indicate
the presence of thymonucleic acid or chromatin in this structure.
tm
93.0 o
70 PROTOZOOLOGY
but fusion of minute vacuoles associated with crescentic Golgi bodies
produces the vacuole.
Duboscq and Grasse who hold that the parabasal body is the
Golgi apparatus, maintain that this body is a source of energy which
is utilized by the motor organellae. Joyet-Lavergne pointed out that
Protozoa
MORPHOLOGY 71
other inclusions, such as the Golgi bodies (in some cases) and certain
bacteria. According to Horning (1926), janus red is said to be a more
exclusive chondriosome stain, as it does not stain bacteria. The
chemical composition of the chondriosome seems to be somewhat
similar to that of the Golgi body; namely, it is a protein compounded
with a lipoidal substance. If the protein is small in amount, it is
resistant to reagents.
The chondriosomes occur as small spherical to oval granules, rod-
filamentous bodies, and show a tendency to adhere to or re-
like or
main near protoplasmic surfaces. In many cases they are distributed
without any definite order; in others, as in Paramecium or Opalina,
they are regularly arranged between the basal granules of cilia
(Horning). In Peranema trichophorum (Fig. 26), according to Hall,
the chondriosomes are said to be located along the spiral striae of
the pellicle. Causey (1925) noticed in Leishmania hrasiliensis usu-
ally eight sphericalchondriosomes in each individual, which become
rod-shaped when the organism divides. He further observed spher-
ical and rod-like chondriosomes in Notiluca scintillans.
72 PROTOZOOLOGY
In certain Protozoa, the chondriosomes are not always demon-
strable. For example, Horning states in Monocystis the chondrio-
somes present throughout the asexual life-cycle as rod-shaped bodies,
but at the beginning of the spore formation they decrease in size and
number, and in the spore none exists. The chondriosomes appear as
soon as the sporozoites are set free. Thus it would appear that the
chondriosomes are reformed de novo. On the other hand, Faure-
Fremiet, the first student of the chondriosomes in Protozoa, main-
tained that they reproduce by division, which has since been con-
firmed by many observers. As a matter of fact. Horning found in
Opalina, the chondriosomes are twisted filamentous structures and
undergo multiple longitudinal fission in asexual division phase. Be-
fore encystment, the chondriosomes divide repeatedly transversely
and become spherical bodies which persist during encystment and
in the gametes. In zygotes, these spherical bodies fuse to produce
longer forms which break up into elongate filamentous structures.
Richardson and Horning further succeeded in bringing about divi-
sion of the chondriosomes in Opalina by changing pH of the medium.
As to the function of chondriosomes, opinions vary. A number of
observers hold that they are concerned with the digestive process.
After studying the relationship between the chondriosomes and
food vacuoles of Amoeba and Paramecium, Horning suggested that
the chondriosomes are the seat of enzyme activity and it is even
probable that they actually give up their own substance for this
purpose. Mast and Doyle hold that the "excretory granules" (chon-
driosomes) in Amoeba proteus contribute to the formation of the
contractile vacuole. The view that the chondriosomes may have
something to do with the cell-respiration expressed by Kingsbury
was further elaborated by Joyet-Lavergne through his studies on
certain Sporozoa. That the chondriosomes are actively concerned
with the development of the gametes of the Metazoa is well known.
Zweibaum's observation, showing an increase in the amount of fatty
acid in Paramecium just prior to conjugation, appears to suggest
this function. On the othr hand. Calkins found that in Uroleptus,
the chondriosomes became abundant in exconjugants, due to trans-
formation of the macronuclear material into the chondriosomes. It
may be stated that the chondriosomes appear to be associated with
the formation of enzymes which participate actively in the processes
of catalysis or synthesis in the protozoan body. The author agrees
with McBride and Hewer who wrote: "it is a remarkable thing that
so little is known positively about one of the 'best known' proto-
plasmic inclusions."
MORPHOLOGY 73
%'
'•^'.*..
%
V
side. The location of the vacuole is not definite in such forms and,
therefore, it moves about with the cytoplasmic movements; and, as
a rule, it is confined to the temporary posterior region of the body.
Although almost spherical in form, it may occasionally be irregular
in shape, as in Amoeba striata (Fig. 161, /). In many testaceans and
heliozoans, the contractile vacuoles which are variable in number,
are formed in the ectoplasm and the body surface bulges out above
the vacuoles at diastole.
In the Mastigophora, the contractile vacuole appears to be more
or less constant in position. In Phytomastigina, they are usually
located in the anterior region and, in Zoomastigina, as a rule, in the
posterior half of the body. The number of the vacuoles present in
an individual varies from one to several.
In the Ciliophora, except Protociliata, there occur one to many
contractile vacuoles, which seem to be located in the deepest part
of the ectoplasm and therefore constant in position. Directly above
each vacuole is found a pore in the pellicle, through which the con-
tent of the vacuole is discharged to outside. In the species of Con-
74 PROTOZOOLOGY
chophthirus, Kidder (1934) observed a narrow slit in the pellicle
on the dorsal surface (Fig. 27). The
just posterior to the vacuole
margin of the slit is thickened and highly refractile. During diastole,
the nearly closed and, at systole, the wall of the contractile
slit is
vacuole appears to break and the slit opens suddenly, the vacuolar
content pouring out slowly. When there is only one contractile
vacuole, it is usually located either near the cytopharynx or, more
often, in the posterior part of the body. When several to many
vacuoles are present, they may be distributed without apparent
order, in linear series, or along the body outline. When the contrac-
tilevacuoles are deeply seated, there is a delicate duct which con-
nects the vacuole with the pore on the pellicle as in Paramecium
per figures are side views; lower figures front views; solid lines indicate
permanent structures; dotted lines temporary structures, a, full diastole;
b-d, stages of systole; e, content of ampulla passing into injection canal;
f, formation of vesicles from injection canals; g, fusion of vesicles to form
content.
In a number of ciliates there occur radiating or collecting canals
besides the main contractile vacuole. These canals radiate from the
central vacuole in Paramecium, Frontonia, Disematostoma, etc. But
when the vacuole is terminal, the collecting canals of course do not
MORPHOLOGY 75
vacuole is in contact (Fig. 28, a). Each vacuole has 5-10 long col-
lecting canals with strongly osmiophilic walls (Fig. 29), and each
canal is made up of terminal portion, a proximal injection canal,
and an ampulla between them. Surrounding the distal portion, there
is osmiophilic cytoplasm which may be granulated or finely reticu-
fibrillarsystems (Fig. 30, d). When the organism divides, the an-
teriordaughter individual retains it, and the posterior individual de-
velopes a new one from the pellicle into which concrement grains
enter after first appearing in the endoplasm. This vacuole shows no
external pore. Dogiel believes that its function is sensory and has
named the vacuole, the statocyst, and the enclosed grains, the
statoliths.
Food vacuoles are conspicuously present in the holozoic Protozoa
which take in whole or parts of other organisms as food. The food
78 PROTOZOOLOGY
vacuole is a space in the cytoplasm, containing the fluid medium
which surrounds the protozoans and which are suspended the
in
food matter, such as various Protophyta, other Protozoa or small
Metazoa. In the Sarcodina, the Mastigophora and the Suctoria,
which do not possess a cytostome, the food vacuoles assume the
shape of the food materials and, when these particles are large, it is
difficult to make out the thin film of water which surrounds them.
When minute food particles are taken through a cytostome, as is
the case with the majority of euciliates, the food vacuoles are usually
spherical and of approximately the same size within a single proto-
zoan. In the saprozoic Protozoa, which absorb fluid substances
through the body surface, food vacuoles containing solid food, of
course,do not occur.
Flagella
Stigma
Pyrenoids
Chromotophores
— Nucleus —
Shell
Chromatophores Pyrenoids
References
Belar, K. 1926 Der Formwechsel der Protistenkerne. Ergebn. u.
Fortschr. Zool., Vol. 6.
Brodsky, a. 1924 Die Trichocysten der Infusorien. Arch. rus.
protist., Vol. 3.
Brown, V. E. 1930 The Golgi apparatus of Amoeba proteus. Biol.
Bull., Vol. 59.
1930 The Golgi apparatus of Pyrsonympha and Dine-
nympha. Arch. f. Protistenk., Vol. 71.
Calkins, G. N. and F. M. Summers (editors). 1941 Protozoa in
biological research. New York.
Causey, D. 1925-1926 Mitochondria and Golgi bodies in Enda-
moeba gingivalis. Mitochondria in Leishniania brasiliensis. Mito-
chondria in Noctiluca scintillans. Univ. Calif. Publ. Zool., Vol.
28.
Chatton, E. and A. Lwoff 1935 Les cili^s apostomes. Arch. zool.
exp. et gen., Vol. 77.
Cleveland, L. R., S. R. Hall, E. P. Sanders and J. Collier 1934
The wood-feeding roach Cryptocercus, its Protozoa, and the
symbiosis between Protozoa and roach. Mem. Amer. Acad. Arts
Sci., Vol. 17.
CusHMAN, J. A. 1933 Foraminifera: their classification and economic
Second edition. Sharon, Mass.
use.
DoFLEiN, F. 1916 Studien zur Naturgeschichte der Protozoen.
VII. Zool. Jahrb. Abt. Anat., Vol. 39.
MORPHOLOGY 81
Physiology
Nutrition
tion," in which the amoeba without contact with the food, forms
84
;
.
PHYSIOLOGY 85
pseudopodia which surround the food on all sides and ingest it (c)
(4) by "invagination," in which the amoeba touches and adheres to
the food, and the ectoplasm in contact with it is invaginated into the
endoplasm as a tube, the cytoplasmic membrane later disappears
(d-h). Jennings, Kepner, Schaeffer and others, have made studies
with reference to the food-ingestion in amoebae.
Fig. 32. Various ways b^^ which amoebae capture food organisms,
a, A moeba verrucosa feeding on Oscillatoria by 'import' (Rhumbler) b, ^
;
nium, etc., where the ciliate attacks other organism and sucks in the
body substance of the latter through the enlarged cytostome.
The ingested food particles are always surrounded by a film of
fluid which envelops the organism and the whole is known as the
food vacuole (p. 77). The quantity taken in with the food
of fluid
varies greatly and, generally speaking, seems to be inversely pro-
it
/T^
g; I
^^ nn
Fig. 34. Ingestion of brine by Rhopalophrya salina (Kirby).
Fig. 35. Diagram showing the digestion within the food vacuoles in
Carchesium polypinurn (Greenwood), a, digestion area; b, region of little
change; c, region of acid reaction; d, region of neutral reaction; e, defeca-
tion area.
very short time. It is generally believed that some substances are se-
creted into the food vacuole by the protoplasm of the organisms, to
stop the activity of the prey within the food vacuole. Engelmann
(1878) demonstrated that the granules of blue litmus, when ingested
by Paramecium or Amoeba, became red in a few minutes. Brandt
PHYSIOLOGY 89
cv
acid during the entire period of protein digestion, and becomes neu-
tral to finally alkaline when the solution of the food substance is
PHYSIOLOGY 91
otic life in the intestine of the termite and of the wood roach, as dem-
onstrated by Cleveland and his co-workers, digest by enzymes the
cellulose which the host insect ingests. The assimilation products
produced by an enormous number of these flagellates are seemingly
sufficient to support the protozoans as well as the host. The cili-
ate commensals inhabiting the stomach of ruminants also appar-
ently digest the cellulose, since the faecal matter as a rule does not
contain this substance. The digestion of fat by Protozoa had not
been known, although oils and fat have been observed in numerous
Protozoa, until Dawson and Belkin (1928) injected different oils
into Amoeha duhia and found that from 1.4 to 8.3 per cent of the
injected oil was digested. Mast (1938) noticed that the neutral fat
92 PROTOZOOLOGY
globules of Colpidium are digested by Amoeba proteus and trans-
formed into fatty acid and glycerine which unite and form neutral
fat.
The indigestible residue of the food is extruded from the body.
The extrusion may take place at any point on the surface in many
Sarcodina by a reverse process of the ingestion of food. But in pelli-
cle-bearing forms, the defecation takes place either through the
cytopyge located body or through an
in the posterior region of the
aperture to the vestibule Carchesium). Permanent cytopyge is
(in
lacking in some forms. In Fdbrea salina, Kirby (1934) noticed that a
large opening is formed at the posterior end, the contents of food
vacuoles are discharged, and the opening closes over. At first the
margin of the body is left uneven, but soon the evenly rounded out-
line is restored. The same seems to be the case with Spirostomum
(Fig. 37), Blepharisma, etc.
hydrates have no direct bearing upon the nutrition, but fatty acids
derived from them participate in the metabolism. Hall, Jahn,
Loefer and others are following the same line of work which may lead
to a better understanding of saprozoic nutrition as found in Proto-
zoa.
The Protozoa which within the body of another organism are
live
able to nourish themselves by absorbing the digested or decomposed
substances of the host and could be considered as saprozoic, though
the term parasitic has sometimes been used. Coelozoic Protozoa be-
long to this group, as for example, Protociliata, astomatous ciliates,
Trypanosomatidae, etc. In the case of cytozoic or certain histozoic
forms, such as Cnidosporidia, the host cytoplasm is apparently
hydrolyzed by enzymes before being absorbed by them.
liquefied or
The which actually feed on host tissue cells, such
parasitic Protozoa,
as Entamoeba histolytica, Balantidium coli, etc., or endocommensals,
employ, of course, the holozoic nutrition.
Many Protozoa nourish themselves by more than one method at
the same or different times, subject to a change in external condi-
94 PROTOZOOLOGY
tions. This is sometimes referred to as mixotrophic nutrition (Pfeif-
fer). For example, Euglena gracilis, according to Zumstein (1900)
and Lwoff (1932) may lose its green coloration and becomes Astasia-
like in the dark, or even in the light when the culture medium is very
abundant in decomposed organic substances, which may indicate
that this organism is capable of carrying on both holophytic and
saprozoic nutrition.
With the introduction of bacteria-free culture technique in recent
years, it has now become well established that a protozoan species
Fig. 38. Form and size variation in Tetrahymena vorax, due to differ-
ences in kind and amount of food material, as seen in life. X400 (Kidder,
Lilly and Claff). a, bacteria-feeder; b, c, saprozoic forms; d, individual
which has fed on killed Colpidium camjnjlum ; e, starved individual from
a killed-Oolpidium culture; f-i, progressive form and size changes of
saprozoic form in the presence of living Colpidium; j, a young carnivore
which has been removed to a culture with living yeast.
96 PROTOZOOLOGY
rate in P. putrinum occurred only in faintly alkaline thyroid ex-
tract inhay infusions. Riddle and Torrey (1923) on the other
hand found that the thja-oxine brought about a slight fall in the
rate of division, an increased rate of pulsation of the contractile
vacuoles, and a decrease in number of excretory crystals in Para-
mecium. The two investigators suggested that the thyroxine acceler-
ates catabolic, and not anabolic, processes. Woodruff and Swingle
(1923, 1924), using a pedigree culture of P. aurelia found that (1)
neither thyroid, pineal, nor pituitary material possesses intrinsic
properties which accelerate division in Paramecium and (2) thyrox-
ine does not accelerate the division and above certain concentrations,
it depresses the division in this ciliate.
100 PROTOZOOLOGY
charide which is insoluble in boiling water, but dissolves in concen-
trated sulphuric acid, potassium hydroxide, and slowly in formalde-
hyde. It does not stain with either iodine or chlor-zinc-iodide and
when treated with a dilute potassium hydroxide, the paramylon
bodies become enlarged and frequently exhibit a concentric stratifi-
cation.
In the Chrysomonadina, the reserve food material is in the form
Protozoa
PHYSIOLOGY 101
Respiration
water entering the body, hence the contractile vacuoles are not
found in them. Just exactly w^hy all euciliates and suctorians possess
the contractile vacuole regardless of habitat, has not fully been ex-
plained. It is assumed that the pellicle of the ciliate is impermeable to
isms were starved, and reappeared when food was given. Schewiakoff
did not see the extrusion of these crystals, but considered that these
crystals were first dissolved and excreted by the contractile vacuoles,
around the vacuoles. In Amoeba proteus,
as they were seen collected
Schubotz (1905) noted that the crystals were of similar chemical
composition and of usually bipyramidal or rhombic form, and that
they measured about 2-5^ in length and doubly refractile. Schaeffer
(1920) observed calcium phosphate cr3^stals in three species of
B
Fig. 41. Examples of crystals present in Protozoa, a-e, in Paramecium
caudatum (Schewiakoff), (a-d, XlOOO, e, X2600); f, in Amoeba proteus;
g, in A. discoides; h-1, in A. duhia (Schaeffer).
Movements
Protozoa move about by means of the pseudopodia, flagella, or
cilia, which may be combined with internal contractile organellae.
held thus that ''the movements of amoebae are due to the presence
of a contractile substance," which was said to be located in the endo-
plasm as a coarse reticulum.'
In the face of advancement of our knowledge on the nature of
protoplasm, Rhumbler realized the difficulties of the surface tension
108 PROTOZOOLOGY
110 PROTOZOOLOGY
lemma at this end (Fig. 45, a), but at other times it is closed by a
thin sheet of gel which prevents the plasmasol from reaching the
anterior end (6). This gel sheet at times persists intact for consider-
able periods, being built up by gelation as rapidly as it is broken
down by stretching, owing to the pressure of the plasmagel against
it.Usually it breaks periodically at various places. Sometimes the
breaks are small and only a few granules of plasmasol pass through
and these gelate immediately and close the openings (d). At other
times the breaks are large and plasmasol streams through, filling the
hyaline cap (c), after which the sol adjoining the plasmalemma gel-
ates forming a new gel sheet. An amoeba is a turgid system, and the
plasmagel isunder continuous tension. The plasmagel is elastic and,
consequently, is pushed out at the region where its elasticity is
weakest and this results in pseudopodial formation. When an amoeba
is elongated and undergoing movement, the elastic strength of the
plasmagel is the highest at its sides, lowest at the anterior end and
intermediate at the posterior end, which results in continuity of the
elongated form and in extension of the anterior end. If pressure is
brought against the anterior end, the direction of streaming of plas-
masol is immediately reversed, and a new hyaline cap is formed at
the posterior end which is thus changed into a new anterior end.
Flagellar movement. The flagellar movement is in a few instances
observable as in Peranema, but in most cases it is very difficult to
observe in life. Since there is difference in the number, location, size,
and probably structure (p. 45) of flagella occurring in Protozoa, it
is supposed that there are varieties of flagellar movements. The first
sure is exerted on the water at right angles to its surface. This pres-
sure can be resolved into two forces: one directed parallel, and the
other at right angles, to the main body axis. The former will drive
the organism forward, while the latter will tend to rotate the animal
on its own axis.
moderate speed, the tip of the flagellum passes through 45° or less
(h-j); (3) when the animal moves backward, the flagellum under-
goes undulation which begins at its base (k-o) (4) when the animal
;
moves to one side, the flagellum becomes bent at right angles to the
body and undulation passes along it from its base to tip (p); and
(5) when the organism undergoes a slight lateral movement, only the
distal end of the flagellum undulates (q).
Ciliary movement. The cilia are the locomotor organella present
permanently in the ciliates and vary in size and distribution among
112 PROTOZOOLOGY
different species. Just as flagellates show various types of move-
ments, so do the ciliates. Individual cilium on a progressing ciliate
bends throughout its length and strikes the water so that the organ-
ism tends to move in a direction opposite to that of the effective
beat, while the water moves in the direction of the beat (Fig. 47,
,,i
/'"^^ 7
1 2
w/'r.i^im^^^ii
Fig. 47. Diagrams illustrating ciliary movements (Verworn). a-d,
movement of a marginal cilium of Urostyla grandis (a, preparatory and
b, effective stroke, resulting in rapid movement; c, preparatory, and d,
effective stroke, bringing about moderate speed) ; e, metachronous move-
ments of cilia in a longitudinal row.
>M:^^M«.
area of a drop of 0.02 per cent acetic acid introduced to the prepara-
tion (Fig. 49, a) ; and if stronger acid is used, the organisms collect
about its periphery where the acid is diluted by the surrounding
water (Fig. 49, h). The reaction to chemical stimuli is probably of
the greatest importance for the existence of Protozoa, since it leads
them to proper food substances, the ingestion of which is the found-
ation of metabolic activities. In the case of parasitic Protozoa,
possibly the reaction to chemical stimuli results in their finding
specifi.c host animals and their distribution in different organs and
tissues within the host body. Recent investigations tend to indicate
that chemotaxis plays an important role in the sexual reproduction
in Protozoa.
PHYSIOLOGY 117
staining dyes. Zuelzer (1905) studied the effects of radium rays upon
various Protozoa and found that the effect was not the same among
different species.For example, limax amoeba was more resistant
than others. In all cases, however, long exposure to the rays was
fatal to Protozoa, the first ejffect of exposure being shown by accele-
rated movement. Halberstaedter and Luntz (1929) studied injuries
and death of Eudorina elegans by exposure to radium rays. Joseph
and Prowazek (1902) found Paramecium and Volvox gave negative
response to the rontgen-ray.
Reaction to temperature stimuli. As was stated before, there
seems to be an optimum temperature range for each protozoan,
118 PROTOZOOLOGY
although it can withstand temperatures which are lower or higher
than that range. As a general rule, the higher the temperature, the
greater the metabolic activities, and the latter condition results in
turn in a more rapid growth and more frequent reproduction. It has
been suggested that change to different phases in the life-cycle of a
protozoan in association with the seasonal change may be largely
due to temperature changes of the environment. In the case of
parasitic Protozoa which pass their life-cycle in two hosts: warm-
blooded and cold-blooded animals, such as Plasmodium and mam-
malian trypanosomes, the change in body temperature of host
animals may bring about specific stages in their development.
Reaction to electrical stimuli. Since Verworn's experiments,
several investigators studied the effects of electric current which
is passed through Protozoa in water. Amoeba shows negative re-
action to the anode andmoves toward the cathode either by revers-
ing the cytoplasmic streaming (Verworn) or by turning around the
body (Jennings). The free-swimming ciliates move mostly toward
the cathode, but a few may take a transverse position (Spirostomum)
or swim to the anode (Paramecium, Stentor, etc.). Of flagellates,
Verworn noticed that Trachelomonas and Peridinium moved to the
cathode, while Chilomonas, Cryptomonas, and Polytomella, swam
to the anode.
References
Ball, G. H. 1925 Studies on Paramecium. I. Uni. Cal. Publ. Zool.,
Vol. 2o.
Becker, E. R. 1941 Effect of parenteral administration of vita-
min Bi and vitamin Be on a Coccidium infection. Proc. Soc.
Exper. Biol. Med., Vol. 46.
and R. I. Dilworth. 1941 Nature of Eimeria nieschulzi
growth-promoting potency of feeding stuffs. II. Vitamins Bi and
Be. Jour. Infect. Dis., Vol. 68.
Berthold, C. 1886 Studien liber Protoplasmamechanik. Leipzig.
Brug, S. L. 1928 Observations on a culture of Entamoeha histoly-
tica. Med. Dienst Volksges. Ned. Indie for 1928.
Calkins, G. N. and F. M. Summers (editors). 1941 Protozoa in
biological research. New York.
Claff, C. L., V. C. Dewey and G. W. Kidder 1941 Feeding
mechanisms and nutrition in three species of Bresslaua. Biol.
Bull., Vol. 81.
Clark, A. M. 1942 Some effects of removing the nucleus from
Amoeba. Australian Jour. Exp. Biol, and Med. Sci., Vol. 20.
Cleveland, L. R. 1925 Toxicity of oxygen for Protozoa in vivo and
in vitro. Animals defaunated without injury. Biol. Bull., Vol. 48.
S. R. Hall, E. P. Sanders and J. Collier 1934
, The
wood-feeding roach Cryptocercus, its Protozoa, and the sym-
biosis between Protozoa and roach. Mem. Amer. Acad. Arts and
Sci., Vol. 17.
PHYSIOLOGY 119
Reproduction
THE modegroups,
reproduction
different
of
although
in Protozoa
it is
is
primarily a
highly variable among
cell division.The
reproduction is initiated by the nuclear division in all cases, which
will therefore be considered first.
Nuclear division
Between a simple hand and a com-
direct division on the one
plicated indirect division which comparable with the typical
is
122
REPRODUCTION 123
ules must be restored sometime before the next division takes place.
Calkins (1926) is "each granule elongates and
of the opinion that
divides into two parts, thus doubling the number of chromomeres."
Reichenow (1928) found that in Chilodonella cucullulus the lightly
Feulgen positive endosome appeared to form chromatin granules
and Kudo (1936) maintained that the large chromatin spherules of
Fig. 51. Division of Endamoeba blattae as seen in life, X250 (Kudo).
The entire process took one hour and seven minutes.
REPRODUCTION 125
't^M^.
and in the absence of this nucleus, the animal perishes. The waste
substances which become accumulated in the macronucleus through
its manifold activities, are apparently eliminated at the time of
a b c d e
Fig. 57. Amitosis of the vegetative nucleus in the trophozoite of
Myxosoma catostonii, X2250 (Kudo).
Stern, very similar to the above. Aside from these two species, the
centriole has been reported in many others, such as Hartmanella
desmose. From the posterior end of each centriole; astral rays extend
out and the spHt chromosomes form loops, pass through "tangled
skein" stage, and emerge as 26 chromosomes. In the metaphase, the
equatorial plate is made up of V-shaped chromosomes as each of the
Fig. 60. Development of spindle and astral rays during the mitosis in
Barbulanympha, X930 (Cleveland), a, interphase centrioles and centro-
somes; b, prophase centrioles with astral rays developing from their distal
ends through the centrosomes; c, meeting of astral rays from two cen-
rays developing into the early central spindle;
trioles; d, astral e, a later
stage showing the entire mitotic figure.
m m
feSsS^iaSt
them soon after division brings forth the long chromosomes still
connected at one end. Thus the chromosomes remain together before
the anaphase begins.
In the instances considered on the preceding pages, the so-called
chromosomes found in them, appear to be essentially similar in
structure and behavior to typical metazoan chromosomes. In many
other cases, the so-called chromosomes or "pseudochromosomes"
are slightly enlarged chromatin granules which differ from the ordin-
ary chromatin granules in their time of appearance and movement
only. In these cases it is of course not possible at present to deter-
mine how and when their division occurs before separating to the
respective division pole. In Table 5 are listed the number of the
"chromosomes" which have been reported by various investigators
in the Protozoa that are mentioned in the present work:
—
REPRODUCTION 139
Protozoa
140
142 PROTOZOOLOGY
of network (c); at this stage, the cytoplasm around the nucleus is
brane still intact (d), which is considered as the end of the prophase.
In the metaphase, the nuclear membrane becomes extremely faint
and the portion over one side of the plate is without it (e). At the
a resting nucleus (h). The two investigators added that if the chro-
matin granules located in the equatorial plate are chromosomes,
"they must be extremely numerous." Liesche (1938) estimates the
number of these granules which he called chromosomes as between
500 and 600.
REPRODUCTION 143
Cytosomic division
Binary fission. As in metazoan cells, the binary fission occurs very
widely among the Protozoa. It is a division of the body through
middle of the extended long axis into two nearly equal daughter
individuals (Fig. 51). In Amocha proteus, Chalkley and Daniel found
that there is a definite correlation between the stages of nuclear divi-
sion and external morphological changes (Fig. 66). During the pro-
b o\SS^
\j^':.
j(..-.^-.
'^-'i5/^:f
'^i^-
r^d ^\,
(Cleveland), etc., the division takes place transversely but the polar-
ity of the posterior individual is reversed so that the posterior end
of the parent organism becomes the anterior end of the posterior
daughter individual. In the ciliate Bursaria, Lund (1917), observed
reversal of polarity in one of the daughter organisms at the time of
division of normal individuals and also in those which regenerated
after being cut into one-half the normal size.
In the Ciliophora the division is as a rule transverse (Fig. 50), in
which the cytosome without any enlargement or elongation divides
by constriction through the middle so that the two daughter indivi-
duals are about half as large at the end of division. Both individuals
usually retain their polarity.
Multiple division. In multiple division the body divides into a
number of daughter individuals, with or without residual cyto-
REPRODUCTION 145
Colony formation
When the division is repeated without a complete separation of
the daughter individuals, a colonial form is produced. The compon-
ent individuals of a colony may either have protoplasmic connections
among them or be grouped within a gelatinous envelope if completely
separated. Or, in the case of loricate or stalked forms, these exo-
skeletal structures may become attached to one another. Although
varied in appearance, the arrangement and relationship of the com-
ponent individuals are constant, and this makes the basis for dis-
tinguishing the types of protozoan colonies, as follows:
146 PROTOZOOLOGY
Catenoid or linear colony. The daughter individuals are attached
endwise, forming a chain of several individuals. It is of compara-
tively rare Examples: Astomatous ciliates such as
occurrence.
Radiophrya Protoradiophrya (Fig. 266) and dinoflagel-
(Fig. 266),
lates such as Ceratium, Haplozoon (Fig. 109) and Polykrikos (Fig.
110).
Arboroid or dendritic colony. The individuals remain connected
with one another in a tree-form. The attachment may be by means
'^^"^^^^^
ml
<^mv--j
Asexual reproduction
The Protozoa nourish themselves by certain methods, grow and
multiply, by the methods described in the preceding pages. This
phase of the life-cycle of a protozoan is the vegetative stage or the
trophozoite. The trophozoite repeats its asexual reproduction process
under favorable circumstances. Generally speaking, the Sporozoa
increase to a much greater number by schizogony and the tropho-
zoites are called schizonts.
Under certain conditions, the trophozoite undergoes encystment
(Fig. 69). Prior to encystment, the trophozoites cease to ingest, and
extrude remains of,food particles, resulting in somewhat smaller
forms which are usually rounded and inactive. This phase is some-
times called the precystic stage. The whole organism becomes de-
differentiated; namely, various cell organs such as cilia, cirri,
flagella, axostyle, peristome, etc., become absorbed. Finally the
organism secretes substances which become solidified into a resistant
wall, and thus the cyst is formed. In this condition, the protozoan
is apparently able to maintain its vitality for a certain length of time
cm ®® ®
Fig. 71. Diagram illustrating the life-cycle of Thelohania legeri (Kudo).
a, extrusion of the polar filament in gut of anopheline larva; b, emerged
amoebula; c-f, schizogony in fat body; g-m, sporont-formation; m-x,
stages in spore-formation.
^')^i-
I
REPRODUCTION 155
-%». *^-.
J''
--7; l-
1 - > '
J - ' '
V V *- «
o ' r
• 4
'> •
;% v«
After a few to several hours, the large masses break down into small
masses (Fig. 78, c) and
conjugants appear in pairs (Fig.
still later,
78, d). The only other ciliate in which mating types are definitely
known to occur is Euplotes patella in which, according to Kimball
(1939), there occurs no agglutination mating reaction.
How widely mating types occur is not known at present. But as
REPRODUCTION 157
was pointed out by Jennings, the mating types may be of general oc-
currence among ciliates; for example, Maupas (1889) observed that
in Lionotus {Loxophyllum) fasciola, Leucophrys patula, Siylonychia
pustulata, and Onychodromus graridis, conjugation took place be-
tween the members of two clones of different origin, and not among
the members of a single clone. Precise information on the occurrence
among different ciliates depends on future research.
In Paramecium aurelia, Sonneborn distinguishes seven varieties
which possess the same morphological characteristics. There occurs
no conjugation between the clones of different varieties. Within
each of the six varieties, there are two mating types, while there is
only one type in the seventh variety. Animals belonging to the
same variety, but to different mating types only conjugate when
put together (Table 6). In P. hursaria, Jennings (1938, 1939) finds
three varieties, but each of two varieties contains four mating types
and in the third variety eight mating types occur (Table 6). In
Euplotes patella, Kimball (1939) observed six mating types (Table
6). These mating types cannot be considered as the true sex types,
since the conjugants mutually fertilize each other.
Recent studies of mating types have revealed much information
regarding conjugation. Conjugation usually does not occur in well-
fed or extremely starved animals, and appears to take place shortly
after the depletion of food. Temperature also plays a role in con-
jugation, as it takes place within a certain range of temperature
which varies even in a single species among different varieties
(Sonneborn). Light seems to have different effects on conjugation
in different varieties of P. aurelia. The time between two conju-
gations also varies in different species and varieties. In P. hursaria,
Jennings found that in some races the second conjugation would
not take place for many months after the first, while in others
such an "immature" period may be only a few weeks. In P. aurelia,
in some varieties there is no "immature" period, while in others there
is 6 to 10 days' "immaturity."
Very little is known about the physiological state of conjugants
as compared with vegetative individuals. Several investigators ob-
served that animals which participate in conjugation show much
viscous body surface. Boell and Woodruff (1941) found that the
mating individuals of Paramecium calkinsi show a lower respiratory
rate than not-mating individuals. Neither is the mechanism of con-
jugation understood at present. Kimball (1942) discovered in
Euplotes patella, the fluid taken from cultures of animals of one
type induces conjugation among the animals of other types. Pre-
—
158 PROTOZOOLOGY
sumably certain substances are secreted by the organisms and be-
come diffused in the culture fluid. In the case of P. aurelia, Sonne-
born considers that there are also some substances which however do
not diffuse into the surrounding medium, and possibly transported
from one individual to another by contact and subsequent migration.
Fuller understanding of the phenomenon of mating types depends
upon future investigations.
When the ciliate possesses more than one micronucleus, the
first division ordinarily occurs in all and the second may or may
Va-
riety
REPRODUCTION 159
Table 6. — Continued
Paramecium hursaria (Jennings)
Variety
160 PROTOZOOLOGY
Nyctotherus cordiformis (Fig. 79) takes place only among those
which live in the tadpolesundergoing metamorphosis (f-j). The
conjugants are said to be much smaller than the ordinary tropho-
synkaryon {g, h) which divides twice into four {i, j). The original
macronucleus undergoes fragmentation and becomes absorbed in the
cytoplasm. Of the four micronuclei, two transform into the new
macronuclei and two remain as micronuclei (k) each dividing into
two after the body divided into two (l).
Another sexual process appears to have been observed by Diller
(1934) in conjugating Paramecium trichium in which there was
no nuclear exchange between the two conjugants. Wichterman
(1939, 1940) observed a similar process in P. caudatum and named it
cytogamy. Two small (about 200m long) individuals of P. caudatum
164 PROTOZOOLOGY
fuse on their oral surfaces. There occur three micronuclear divisions
as in the case of conjugation, but there no nuclear exchange be-
is
:^I
^sS^?-^
^U
gelatinous envelope. Both nuclei divide twice and produce four nu-
clei, The two daughter cells, each with one
three of which degenerate.
haploid nucleus, undergo paedogamy and the resulting individual
now contains a diploid nucleus.
In Paramecium aurelia, Diller (1936) found simple fragmentation
of the macronucleus which was not correlated with any special
micronuclear activity and which could not be stages in conjugation
or autogamy. Diller suggests that if conjugation or autogamy is to
create a new nuclear complex, as is generally held, it is conceivable
that somewhat the same result might be achieved by 'purification
act'(through fragmentation) on the part of the macronucleus itself,
without involving micronuclei. He coined the term hemixis to in-
clude these reorganizations.
which germ and soma cells are differentiated. Since that time, the
problem of potential immortality of Protozoa has been a matter
which attracted the attention of numerous investigators. Because of
large dimensions, rapid growth and reproduction, and ease with
which they can be cultivated in the laboratory, the majority of
Protozoa used in the study of the problem have been free-living
freshwater ciliates that feed on bacteria and other microorganisms.
The very first extended study was made by Maupas (1888) who
isolated Stylonychia pustulata on Februar}^ 27, 1886, and observed
316 binary fissions until July 10. During this period, there was noted
a gradual decrease in size and increasing abnormality in form and
structure, until the animals could no longer divide and died (Fig.
84). A large number of isolation culture experiments have since been
carried on numerous species of ciliates by many investigators. The
results obtained are not in agreement. However, the bulk of ob-
tained data indicates that the vitality of animals decreases with the
passing of generations until finally the organisms suffer inevitable
death, and that in the species in which conjugation or other sexual
reproduction occurs, the declining vitality becomes restored. Perhaps
the most thorough experiment was carried on by Calkins (1919,
1933) with Uroleptus mohilis. Starting with an exconjugant on
168 PROTOZOOLOGY
November 17, 1917, a series of pure-line cultures was established by
the daily isolation method. It was found that no series lived longer
than a year, but when two of the progeny of a series were allowed to
conjugate after the first 75 generations, the exconjugants repeated
the history of the parent series, and did not die when the parent
series died. In this way, lines of the same organism have lived for
more than 12 years, passing through numerous series. In a series,
the average division for the first 60 days was 15.4 divisions per 10
days, but the rate gradually declined until death. Woodruff and
Spencer (1924) also found the isolation cultures of Spathidium
spatula (fed on Colpidium colpoda) died after a gradual decline in
the division rate, but were inclined to think that improper environ-
mental conditions rather than internal factors were responsible for
the decline.
On the other hand. Woodruff (1932) found that 5071 generations
produced by binary fission from a single individual of Paramecium
aurelia between May 1, 1907 and May 1, 1915, did not manifest
any decrease in vitality after eight years of uninterrupted asexual
reproduction without conjugation. With a race of P. caudatum,
Metalnikov (1924) observed a similar continued asexual reproduc-
tion. Dawson (1919) subjected an amicronucleate race of Oxytricha
hymenostoma to isolation culture and found that it declined in divi-
sion-rate and finally died out; but in mass cultures, the organisms
lived indefinitely. He attributed the decline in isolation culture to
improper environmental conditions. With Actinophrys sol Belaf
(1924) carried on isolation cultures (thus preventing paedogamy (p.
164) for 1244 generations for a period of 32 months and noticed no
decline in the division rate. Hartmann (1921) made a similar obser-
vation on Eudorina elegans. It would appear that in these forms the
life continues indefinitely without apparent decrease in vital activity.
As has been noted in the beginning part of the chapter, the
macronucleus in the ciliates undergoes, at the time of binary fission
a reorganization process before dividing into two parts and undoubt-
edly, there occurs at the same time extensive cytoplasmic reorgani-
zation as judged by the degeneration and absorption of the old, and
formation of the new, organellae. It is reasonable to suppose that
this reorganization of the whole body structure at the time of divi-
sion is an elimination process of waste material accumulated by the
organism during the various phases of vital activities as was con-
sidered by Kidder and others (p. 127) and that this elimination,
though not complete, enables the protoplasm of the products of divi-
sion to carry on their metabolic functions more actively.
REPRODUCTION 169
to their progeny, but that of older stocks results in many strains with
restored vitality. Sonneborn (1942) mentions the appearance of
varied characters after macronuclear regeneration (p. 129) in P.
aurdia. The animals become smaller and of different form; they
reproduce more slowly they are less viable and die out after a period
;
Regeneration
The capacity of regenerating the lost parts, though variable
among different species, is characteristic of all Protozoa from simple
forms to those with highly complex organizations, as shown by ob-
servations of numerous Brandt (1877) studied regen-
investigators.
eration in Actinosphaerium eichhorni and found that only nucleate
portions containing at least one nucleus regenerated, and enucleate
portions or isolated nuclei degenerated. Similarly Gruber (1886) found
in Amoeba proteus the nucleate portion regenerated completely,
while enucleate part became rounded and perished in a few days.
The parts which do not contain nuclear material, may continue to
show certain metabolic activities such as locomotion, contraction of
contractile vacuoles, etc., for some time; for example, Grosse-Aller-
mann (1909) saw enucleate portions of Amoeba verrucosa alive for
20 to 25 days, while Stole (1910) found enucleate Amoeba proteus
living for 30 days. Clark (1942, 1943) showed that Amoeba proteus
lives for about seven days after it has been deprived of its nucleus.
Enucleated individuals show a 70 per cent depression of respiration
and are unable to digest food due to the failure of zymogens to be
activated in the dedifferentiating cytoplasm. It is now a well estab-
lished fact that when a protozoan is cut into two parts and the parts
are kept under proper environmental conditions, the enucleated
portion is able to carry on catabolic activities, but unable to under-
take anabolic activities, and consequently degenerates sooner or
later.
In Arcella (Martini; Hegner) and Difflugia (Verworn; Penard),
when the tests are partially destroyed, the broken tests remain un-
REPRODUCTION 171
at 51°C.
Such differences or varieties appear to be due to the influence of
diverse environmental conditions, and will continue to existunder
these conditions; but when the organisms of different varieties are
subjected to a similar environment, the strain differences disappear
sooner or later. That the differences in kind and amount of foods
bring about extremely diverse individuals in Tetrahymena vorax and
Chilomonas Paramecium in bacteria-free cultures has already been
mentioned(p. 94). Chlamydomonas dcbaryana are represented by
many races differing in form, size, and structure, in various localities
as well as under different laboratory conditions. Moewus (1934) dis-
176
VARIATION AND HEREDITY 177
races. It was found that the longer a strain has remained under con-
ditions producing a given type, the greater the time and the number
of generations needed to change it to a new type under a new condi-
tion, as is shown in Table 7.
While in many species, the races or varieties have apparently been
brought about into being under the influence of environmental con-
ditions, in others the inherited characters persist for a long period,
and still in others the biotype may show different inherited char-
—
178 PROTOZOOLOGY
acters. To the last-mentioned category belongs perhaps a strain of
Tetrahymena geleii in which, according to Fiirgason (1940), a pure-
line bacteria-free culture derived from a single individual was found
to becomposed of individuals differing in shape and size which be-
came more marked in older cultures.
The first comprehensive study dealing with the variation in size
and its inheritance in uniparental or vegetative reproduction of
into the same host. Hoare (1943) reviewed recently the "biological"
races of certain parasitic Protozoa.
Jollos (1921) subjected Paramecium caudatum to various environ-
180 PROTOZOOLOGY
mental influences such as temperature and chemicals, and found that
the animals develop tolerance which is inherited through many gen-
erations even after removal to the original environment. For exam-
ple, one of the clones which tolerated only 1.1% of standard solution
of arsenic acid, was cultivated in gradually increasing concentrations
for four months, at the end which the tolerance for this chemical
of
was raised to 5%. After being removed to water without arsenic
acid, the tolerance changed as follows: 22 days, 5%; 46 days, 4.5%;
151 days, 4%; 166 days, 3%; 183 days, 2.5%; 198 days, 1.25% and
255 days, 1%. As the organisms reproduced about once a day, the
acquired increased tolerance to arsenic was inherited for about 250
generations.
There are also known inherited changes in form and structure
which are produced under the influence of certain environmental
conditions. Jollos designated these changes long-lasting modifica-
tions (Dauermodifikationen) and maintained that a change in en-
vironmental conditions, if applied gradually, brings about a change,
not in the nucleus, but in the cytoplasm, of the organism which
when transferred to the original environment, is inherited for a
number of generations. These modifications are lost usually during
sexual processes at which time the whole organism is reorganized.
The long-lasting morphological and physiological modifications
induced by chemical substances have long been known in parasitic
Protozoa. Werbitzki (1910) discovered that Trypanosoma hrucei
loses its blepharoplast when inoculated into mice which have
been treated with pyronin, acridin, oxazin and allied dyes. Laveran
and Roudsky (1911) found that these dyes have a special affinity for,
and bring about the destruction by auto-oxidation of, the blepharo-
plast.Such trypanosomes lacking blepharoplast behave normally
and remain in that condition during many passages through mice.
When subjected to small doses of certain drugs repeatedly, species
of Trypanosoma often develop into drug-fast or drug-resistant
strains which resist doses of the drug greater than those used for the
treatment of the disease for which they are responsible. These modi-
fications may also persist for several hundred passages through host
animals and invertebrate vectors, but are eventually lost.
Long-lasting modifications have also been produced by several
investigators by subjecting Protozoa to various environmental in-
fluences during the nuclear reorganization at the time of fission,
conjugation, or autogamy. In Stentor (Popoff) and Glaucoma
(Chatton), long-lasting modifications appeared during asexual divi-
sions. Calkins (1924) observed a double-type Uroleptus mohilis
VARIATION AND HEREDITY 181
P. uvella
Form A: Oval (F), without papilla (p), with stigma (S), large (D)
(Fig. 87, a).
Form B: Oval (F), without papilla (p), without stigma (s), large (D)
(Fig. 87, 6).
P. pascheri
Form C: Pyriform (f), with papilla (P), without stigma (s), large
(D)(Fig.87,c).
Form D: Pyriform (f), with papilla (P), without stigma (s), small
(d)(Fig.87,6;).
VARIATION AND HEREDITY 183
of course found. It was found that about half the zygotes gives rise to
the two parental combinations (Fig. 87, h, c), while the other half
gives rise to FPsD (Fig. 87, e) and fpsD (Fig. 87,/).
When B (FpsD) is crossed with D (fPsd) or A (FpSD) is crossed
with D (fPsd), only two types of swarmers are also formed from
each zygote, and in the case of BXD, eight different combinations
are produced, while in the case of AXD, sixteen different combina-
tions, which appear in about equal numbers, are formed. Thus these
four factors or characters show independent assortment during divi-
sions of the zygote.
Furthermore, Moewus noticed that certain other characters ap-
peared to be linked with some of the four characters mentioned
above. For example, the length of flagella, if it is under control of a
factor, is linked on the same chromosome with the size-controlling
factors (D, d), for large individuals have invariably long flagella
and small individuals short flagella. During the experiments to de-
termine this linkage, it was found that crossing over occurs between
two entire chromosomes that are undergoing synapsis.
184 PROTOZOOLOGY
In certain races of Polytoma pascheri and Chlamydomonas euga-
metos, the sexual fusion takes place between members of different
clones only. The zygote gives was stated before to four swarm-
rise as
ers by two divisions, which are evenly divided between the two
sexes, which shows that the sex-determining factors are lodged in a
single chromosome pair. In a cross between Chlamydomonas para-
doxa and C. pseudoparadoxa, both of which produce only one type of
gamete in a clone, the majority of the zygotes yield four clones, two
producing male gametes and the other two female gametes; but a
small number of zygotes gives rise to four clones which contain both
gametes. It is considered that this is due to crossing-over that
brought the two sex factors (P and M) together into one chromo-
some, and hence the "mixed" condition, while the other chromosome
which is devoid of the sex factors gives rise to individuals that soon
perish.
In crosses between Chlamydomonas eugametos which possesses a
stigma and 10 haploid chromosomes and C. paupera which lacks a
stigma and 10 haploid chromosomes, 12 pairs of factors including
sex factor are distinguishable. Consequently at least two chromo-
somes must have two factors in them. Thus adaptation to acid or
alkaline culture media was found to be linked with differences in
the number of divisions in zygote. That there occurs a sex-linked in-
heritance in Chlamydomonas was demonstrated by crossing stigma-
bearing C. eugametos of one sex with stigma-lacking C. paupera of
the opposite sex. The progeny that were of the same sex as C. euga-
metos parent possessed stigma, while those that were of the same sex
as C. paupera parent lacked stigma. Thus it is seen that the sex factor
and stigma factor are located in the same chromosome.
The genetics of conjugation which takes place between two diploid
conjugants has been studied by various investigators. Pure-line
cultures of exconjugants show that conjugation brings about diverse
inherited constitutions in the clones characterized by difference in
size, form, division-rate, mortality-rate, vigor, resistance, degene-
ration, etc. The diversities brought about by autogamy are not as
varied as those produced by conjugation. In Paramecium much in-
formation has in recent years been brought to light through the
studies of Sonneborn, Jennings, Kimball, and others on the mating
type (p. 156). The mating type is as a rule inherited without change to
descendants through vegetative reproduction.
Sonneborn (1939) has made extended studies of variety 1 of
Paramecium aurelia (p. 158) and found that genetically diverse ma-
terials show different types of inheritance, as follows:
VARIATION AND HEREDITY . 185
sion, one and all its progeny, are of one mating type, and the other
and all its progeny are of the other mating type. A similar change
was also found to take place at autogamy. Sonneborn considers
that the mating types are determined by macronuclei, as judged by
segregation at first or sometimes second division in exconjugants and
by the influence of temperature during conjugation and the first
division.
(2) Stocks containing only one mating type. No conjugation oc-
curs in such stocks. Autogamy does not produce any change in type
which is always type I. Stocks that contain type II only have not
yet been found.
(3) Hybrids between stocks containing one and two mating types.
When the members of the stock containing both types I and II
(two-type condition) conjugate with those of the stock containing
one type (one-type condition), all the descendants of the hybrid
exconjugants show two-type condition, which shows the dominancy
of two-type condition over one-type condition. The factor for the
two-type condition may be designated A and that for the one-type
condition a. The parent stocks are AA and aa, and all Fi hybrids Aa.
When the hybrids (Aa) are backcrossed to recessive parent (aa)
(158 conjugating pairs in one experiment), approximately one-half
(81) of the pairs give rise to two-type condition (Aa) and the remain-
ing one-half (77) of the pairs to one-type condition (aa), thus showing
a typical Mendelian result. When Fi hybrids (Aa) were interbred by
120 conjugating pairs, each exconjugant in 88 of the pairs gave
rise to two-type condition and each exconjugant in 32 pairs pro-
duced one-t3^pe condition, thus approximating an expected Men-
delian ratio of 3 dominants to 1 recessive. That the F2 dominants
are composed of two-thirds heterozygotes (Aa) and one-third homo-
zygotes (AA) was confirmed by the results obtained by allowing F2
dominants to conjugate with the recessive parent stock (aa). Of 19
pairs of conjugants, 6 pairs gave rise to only dominant progeny,
which shows that they were homoz^^gous (AA) and their progeny
heterozygous (Aa), while 13 pairs produced one-half dominants and
one-half recessives, which indicates that they were heterozygous
(Aa) and their progeny half homozygous (aa) and half heterozygous
186 .
PROTOZOOLOGY
(Aa). Thus the genie agreement between two conjugants of a pair
and the relative frequency of various gene combinations as shown in
these experiments confirm definitely the occurrence of meiosis and
chromosomal exchange during conjugation which have hitherto been
known only on cytological ground.
In Euplotes Kimball (1942) found that the six mating
patella,
types (p. 159) are determinedby six possible combinations of a series
of three allelic genes. There is no dominance among these alleles, the
three heterozygous combinations determining three mating types
being different from one another and from the three determined by
homozygous combination. Kimball (1939, 1941) had shown that
the fluid obtained free of Euplotes from a culture of one mating type
will induce conjugation among animals of certain other mating
types. When all possible combinations of fluids and animals are made
it was found that the fluid from any of the heterozygous types in-
duces conjugation among animals of any types other than its own
and the fluid from any of the homozygous types induces conjugation
only among animals of the types which do not have the same allele
as the type from which the fluid came. These reactions are explained
by an assumption that each of the mating type alleles is responsible
for the production by the animal of a specific conjugation-inducing
substance. Thus the two alleles in a heterozygote act independently
of each other; each brings about the production by the animal of a
substance of its own. Thus heterozygous animals are induced to con-
jugate only by the fluids from individuals which possess an allele
not present in the heterozygotes.
The relation between the cytoplasm and nucleus in respect to in-
heritance has become better known in recent years in some ciliates.
References
Calkins, G. N. 1924 Uroleptus mohilis. V. Jour. Exp. Zool., Vol.
41.
De Garis, C. F. 1935 Heritable effects of conjugation between free
individuals and double monsters in diverse races of Paramecium.
Ibid., Vol. 71.
FuRGASON, W. H. 1940 The significant cytostomal pattern of the
"Glaucoma-Colpidium group," and a proposed new genus and
species, Tetrahymena
geleii. Arch. f. Protistenk., Vol. 94.
Hegner, R. W. 1919 Heredity, variation, and the appearance of
diversities during the vegetative reproduction of Arcella dentata
Genetics, Vol. 4.
HoARE, C, A. 1943 Biological races in parasitic Protozoa. Biol. Re-
views, Vol. 18.
Jennings, H. S. 1909 Heredity and variation in the simplest or-
ganisms. Amer. Nat., Vol. 43.
1916 Heredity, variation and the results of selection in the
uniparental reproduction of Difflugia corona. Genetics, Vol. 1.
1929 Genetics of the Protozoa. BibUographia Genetica,
Vol. 5.
1937 Formation, inheritance and variation of the teeth in
Difflugia corona. Jour. Exp. Zool., Vol. 77.
1939 Genetics of Paramecium hursaria. I. Mating types and
groups, their interrelations and distribution; mating behavior
and self-sterility. Genetics, Vol. 24.
1941 Inheritance in Protozoa. In G. N. Calkins and F. M.
Summers (editors) Protozoa in biological research. New York.
:
VARIATION AND HEREDITY 189
References
BuTSCHLi, O. 1883-1887 Bronn's Klassen und Ordnungen des
Thierreichs. Vol 1.
DoFLEiN, F. and E. Reichenow 1929 Lehrhuch der Protozoenkunde.
Jena.
MiNCHiN, E. A. 1912 Introduction to the study of the Protozoa. Lon-
don.
Pascher, a. 1912 Ueber Rhizopoden- und Palmellastadien bei
Flagellaten. Arch. f. Protistenk.,, Vol. 25.
1917 Rhizopodialnetz als Fangvorrichtung bei einer Plas-
modialen Chrysomonade. Ibid., Vol. 37.
1917 Fusionsplasmodien bei Flagellaten und ihre Bedeut-
ung fiir die Ableitung der Rhizopoden von den Flagellaten.
Ibid.
1917 Flagellaten und Rhizopoden in ihren gegenseitigen
Beziehungen. Ibid., Vol. 38.
ScHERFFEL, A. 1901 Kleiner Beitrag zur Phylogenie einiger Grup-
pen niederer Organismen. Bot. Zeit., Vol. 59.
ZuMSTEiN, H. 1900 Zur Morphologic von Euglena gracilis Klebs.
Pringsheims Jahrb., Vol. 34.
Chapter 8
200 PROTOZOOLOGY
mal. Trypanosoma, a representative genus of the latter group, in-
cludes important disease-causing parasites of man and of domestic
animals.
The Mastigophora are divided into two subclasses as follows
(^^
calities (Ahlstrom).
Genus Hyalobryon Lauterborn. Solitary or colonial; individual
body structure similar to that of Dinobryon; lorica in some cases
tubular, and those of 'young individuals are attached to the exterior
of parent tests; fresh water.
H. ramosum L. (Fig. 92, g). Lorica 50-70iu long by 5-9iu in diame-
ter; body up to 30m by 5ix; on vegetation in standing fresh water.
Genus Stylopyxis Bolochonzew. Solitary; body located at bottom
of a delicate stalked lorica with a wide aperture; 2 lateral chromato-
phores; fresh water.
S. mucicola B. (Fig. 92, h). Lorica 17-18m long; stalk about 33/^
long; body 9-1 Iju long: fresh water.
References
BtJTSCHLi, O. 1883-1887 Mastigophora. In: Bronn's Klassen und
Ordnungen des Thierreichs. Vol. 1, part 2.
DoFLEiN, F. and E. Reichenow. 1929 Lehrhuch der Protozoen-
kunde. Jena.
Kent, S. 1880-1882 A manual of Infusoria. London.
Pascher, A. 1914 Die Siisswas-
Flagellatae: Allgemeiner Teil. In:
Part 1.
serflora Deutschlands.
Stein, F. 1878, 1883 Der Organismus der Infusionsthiere. 3 Abt.
Der Organismus der Flagellate oder Geisselinfusorien. Parts 1 and
2. Leipzig.
THE cryptomonads
constant body form.
differ from the chrysomonads in having a
Pseudopodia are very rarely formed, as
the body covered by a pelHcle. The majority show dorso-ventral
is
tractile vacuoles, one to several, are simple and are situated near the
cj^topharynx. A single vesicular nucleus is ordinarily located near
the middle of the body.
Asexual reproduction, by longitudinal fission, takes place in
either the active or the non-motile stage. Sexual reproduction is un-
known. Some cryptomonads form palmella stage and others gelati-
nous aggregates. In the suborder Phaeocapsina, the palmella stage is
213
214 PROTOZOOLOGY
Family 1 Cryptomonadidae Stein
Genus Cryptomonas Ehrenberg. Body elliptical with a firm pel-
licle; anterior end truncate; dorsal side convex, ventral side slightly
so or flat; nucleus posterior; longitudinal furrow; tubular cavity
extending to the middle of body, through which equally long flagella
arise; 2 lateral chromatophores vary in color from green to blue-
green, brown or rarely red; holophytic; with small starch-like bodies
R. lens Pascher and Ruttner (Fig. 95, g). Spindle-form; about 16m
long; in fresh water.
fresh water.
216 PROTOZOOLOGY
P. phaeophycearum P. (Fig. 95, i). 15-17)u by 7-9 fi.
References
Feitsch, F. E. 1935 The structure and reproduction of the algae.
Cambridge.
Pascher, A. 1913 Cryptomonadinae. Susswasserflora Deutschlands,
etc. part 2. Jena.
West, G. S, and F. E. Fritsch. 1927 A treatise on the British _
THE phytomonads
with a
lates,
are
resemblance
close
more
the
small,
to
or less rounded, green flagel-
algae. They show a definite
body form, and most of them possess a cellulose membrane, which
is thick in some and thin in others. There is a distinct opening in the
217
218 PROTOZOOLOGY
reproduction and palmella formation known; sexual reproduction
isogamy or anisogamy fresh water. Numerous species.
;
Family 2 Trichlorididae
With three flagella.
Genus Trichloris Scherffel and Pascher. Bean-shape; flagellate
side flattened or concave; opposite side convex; chromatophore
large, covering convex side; 2 pyrenoids surrounded by starch
granules; a stigma near posterior end of chromatophore; nucleus
central; numerous contractile vacuoles scattered; 3 flagella near
anterior end.
T. paradoxa S and P. (Fig. 97, j). 12-15m broad by 10-12^ high;
flagella up to 30m long.
Family 3 Carteriidae
With four flagella arising from anterior pole.
Genus Carteria Diesing {Corhierea, Pithiscus Dangeard; Tetra-
mastix Korschikoff). Ovoid, chromatophore cup-shaped; pyrenoid;
stigma; 2 contractile vacuoles; fresh water. Numerous species.
C. cordiformis (Carter) (Fig. 98, a). Heart-shaped in front view;
ovoid in profile; chromatophore large; 18-23m by 16-20^.
C. ellipsoidalis Bold. Ellipsoid; chromatophore; a small stigma;
division into 2, 4, or 8 individuals in encysted stage; 6-24m long;
fresh water, Maryland (Bold, 1938).
Genus Pyramimonas Schmarda (Pyramidomonas Stein). Small
pyramidal or heart-shaped body; with bluntly drawn-out posterior
end; usually 4 ridges in anterior region; 4 flagella; green chromato-
PHYTOMONADINA 223
Family 4 Chlorasteridae
With 5 flagella arising from anterior pole.
Genus Chloraster Ehrenberg. Similar to Pyramimonas, but an-
terior half with a conical envelope drawn out at four corners; with 5
flagella fresh or salt water.
;
C. gyrans E. (Fig. 98, h). Up to 18m long; standing water; also re-
ported from salt water.
within the shell; valves may become separated from each other ow-
ing to an increase in gelatinous contents.
Genus Phacotus Perty. Oval to circular in front view; lenticular
in profile; protoplasmic body does not fill dark-colored shell com-
pletely; flagella protrude through a foramen; asexual reproduction
into 2 to 8 individuals fresh water.
;
S. pluvialis C. (Figs. 74; 100, e). Cells 7-13m long; colony 30-60/i
in diameter.
Genus Pandorina Bory. Spherical or subspherical colony of usu-
ally 16 (sometimes 8 or 32) biflagellate individuals, closely packed
230 PROTOZOOLOGY
within a gelatinous, but firm and thick matrix; individuals often
angular; with stigma and chromatophores; asexual reproduction
through simultaneous division of component individuals; anisogamy
preceded by division of each cell into 16 to 32 gametes; zygotes
colored and covered by a smooth wall; fresh water. One species.
P. morum (Miiller) (Figs. 75; 100, /). Cells 8-17m long; colony
20-40/x, up to 250/i in diameter; ponds and ditches.
Genus Mastigosphaera Schewiakoff. Similar to Pandorina; but
individuals with a single flagellum which is 3.5 times the body length;
fresh water.
M. gohii S. (Fig. 100, g). Individual 9m long; colony 30-33^.
Genus Eudorina Ehrenberg. Spherical or ellipsoidal colony of
usually 32 or sometimes 16 spherical cells; asexual reproduction
similar to that of Pandorina; sexual reproduction with 32-64 spheri-
cal green macrogametes and numerous clustered microgametes; red-
dish zygote with a smooth wall; fresh water.
E. elegans E. (Figs. 76; 100, h). Cells 10-24^ in diameter; colony
40-1 50^1 in diameter; in ponds, ditches and lakes.
Genus Pleodorina Shaw. Somewhat similar to Eudorina, being
composed of 32, 64, or 128 ovoid or spherical cells of 2 types: small
somatic and large generative, located within a gelatinous matrix;
fresh water.
P. illinoisensis Kofoid (Figs. 31, h, c; 100, i). 32 cells in ellipsoid
References
THE bodybody
definite
is some are plastic, others have a
as a rule elongated;
form with a well-developed, striated or variously
sculptured pellicle. At the anterior end, there is an opening through
which a flagellum protrudes. In holophytic forms the so-called cyto-
stome and cytopharynx, if present, are apparently not concerned with
the food-taking, but seem to give a passage-way for the flagellum
and also to excrete the waste fluid matters which become collected
in one or more contractile vacuoles located around the reservoir.
In holozoic forms, a well-developed cytostome and cytopharynx are
present. Ordinarily there is only one flagellum, but some possess two
or three. Chromatophores are present in the majority of the Eu-
glenidae, but absent in two families. They are green, vary in
shape, such as spheroidal, band-form, cup-form, discoidal, or
fusiform,and usually possess pyrenoids. Some forms may contain
haematochrome. A small but conspicuous stigma is invariably pres-
ent near the anterior end of the body in chromatophore-bearing
forms.
Reserve food material is the paramylon body, fat, and oil, the
presence of which depends naturally on the metabolic condition
of the organism. The paramylon body assumes diverse forms in dif-
ferent species, but is, as a rule, constant in each species, and this
facilitates specific identification to a certain extent. Nutrition is
232
;
EUGLENOIDINA 233
stalk; body ovoidal, plastic; attached to test with its posterior end;
a single flagellum; a stigma; numerous chromatophores discoid;
with or without pyrenoids; reproduction as in Trachelomonas
fresh water.
A. vaginicola S. (Fig, 102, n). Lorica about 43;u b}^ 15ju.
Genus Colacium Ehrenberg. Stalked individuals form colony;
frequently attached to animals such as copepods, rotifers, etc; stalk
mucilaginous; individual cells pyriform, ellipsoidal or cylindrical;
without flagellum; a single flagellum only in free-swimming stage;
discoidal chromatophores numerous; with pyrenoids; multiplication
by longitudinal fission; also by swarmers, possessing a flagellum and
a stigma; fresh water. Several species.
238 PROTOZOOLOGY
C. vesiculosum E, (Fig. 102, o). Colony of 2-8 cells; also solitary;
scribed by Stein.
C. subtilis D. (Fig. 73). 7-20^ long.
M. picta Faria, da Cunha and Pinto (Fig. 103, s). In salt water;
Rio de Janeiro.
References
Allegre, C. F. and T. L. Jahn 1943 A survey of the genus
Phacus Dujardin (Protozoa; Euglenoidina). Trans. Amer. Micr.
Soc, Vol. 62.
Dangeard, p. 1901 Recherches sur les Eugl^niens. Le Botaniste.
P. 97.
Fritsch, F. E. 1935 The structure and reproduction of the algae.
Vol. 1. Cambridge.
Hall, R. P. 1923 Morphology and binary fission of Menoidium
incurvum (Fres.) Klebs. Uni. Cal. Publ. Zool., Vol. 20.
Johnson, L. P. 1939 A
study of Euglena rubra Hardy 1911. Trans.
Amer. Micr. Soc, Vol. 58.
1944 Euglenae of Iowa. Ibid. Vol. 63.
and T. L. Jahn 1942 Cause of the green-red color change in
Euglena rubra. Physiol. Zool., Vol. 15.
Lemmermann, E. 1913 Eugleninae. In: Siisswasserfl. Deutschlands,
Part 2.
Pascher, a. 1913 Chloromonadinae. Ibid. Part 2.
Smith, G. M. 1933 The freshwater algae of the United States. New
York.
Wenrich, D. H. 1924 Studies on Euglenamorpha hegneri n. g.,
n. sp., a euglenoid flagellate found in tadpoles. Biol. Bull., Vol.
47.
West, G. S. and F. E. Fritsch. 1927 A treatise on the British fresh-
water algae. Cambridge.
Chapter 12
Annulus or girdle
Hypocone
245
246 PROTOZOOLOGY
in a circle near the anterior end. The longitudinal flagellum often
projects beyond the body and vibrates. Combination of the move-
ments of these flagella produces whirHng movements characteristic
of the organisms.
The majority of dinofiagellates possess a single somewhat massive
nucleus with evenly scattered chromatin, and usually several endo-
somes. There are two kinds of vacuoles. One is often surrounded by
a ring of smaller vacuoles, while the other is large, contains pink-
colored fluid and connected with the exterior by a canal opening into
a flagellar pore. The latter is known
as the pusule which functions
as a digestive organella (Kofoid and Swezy). In many freshwater
forms a stigma is present, and in Pouchetiidae there is an ocellus
composed of an amyloid lens and a dark pigment-ball. The majority
of planktonic forms possess a large number of small chromatophores
which are usually dark yellow, brown or sometimes slightly greenish
and are located in the periphery of the body, while bottom-dwelling
and parasitic forms are, as a rule, colorless, because of the absence of
chromatophores. A few forms contain haematochrome. The method
of nutrition is holophytic, holozoic, saprozoic, or mixotrophic. In
holophytic forms, anabolic products are starch, oil, or fats.
Asexual reproduction is bj^ binary or multiple fission or budding
in either the active or the resting stage and differs among different
groups. Encystment is of common occurrence. In some forms the
cyst wall formed within the test. The cysts remain alive for many
is
years; for example, Ceratium cysts w^ere found to retain their vital-
ity in one instance for six and one-half years. Conjugation and sexual
fusion have been reported in certain forms, but definite knowledge on
sexual reproduction awaits further investigation.
The dinofiagellates are abundantplankton of the sea and
in the
play an important part in the economy of marine life as a whole. A
number of parasitic forms are also known. Their hosts include vari-
ous diatoms, copepods and several pelagic animals.
Bivalve shell without furrows Suborder 1 Prorocentrinea
Naked or with shell showing furrows. .Suborder 2 Peridiniinea (p. 248)
Naked; without furrows; no transverse flagellum
Suborder 3 Cystoflagellata (p. 261)
Fig.
107. a, Pronoctiluca tentaculatum, X730 (Kofoid and Swezy);
b, Oxyrrhis marina, X840 (Senn); c. Pouchetia fusus, X340 (Schiitt);
d, P. maxima, X330 (Kofoid and Swezy); e, Protopsis ochrea, X340
(Wright); f, Nematodinium partitum, X560 (Kofoid and Swezy); Pro- g,
tenjthropsis crassicaudata, x740 (Kofoid and Swezy); h, Erythropsis
cormita, X340 (Kofoid and Swezy); i,
j , Nodiluca scintillans (i, side view;
j, budding process), Xl40 (Robin),
DINOFLAGELLATA 251
Numerous species.
P. tahulatum Claparede and Lachmann (Fig. 110, c). 48m by 44^;
fresh water.
References
Chatton, E. 1920 Les P^ridieniens parasites morphologie, repro-
;
water.
M. lacustris Lauterborn (Fig. 113, 6). Multinucleate; 30-40/ii in
diameter; fresh water.
Family 2 Mastigamoebidae
With 1-3 or and axopodia or lobo podia; uninucle-
rarely 4 flagella
ate; flagellum from a basal granule which is connected
arises
with the nucleus by a rhizoplast; binary fission in both trophic and
encysted stages; sexual reproduction has been reported in one spe-
263
264 PROTOZOOLOGY
cies; holozoic or saprozoic; the majority are free-living, though a few
parasitic.
Genus Mastigamoeba Schulze {Mastigina Frenzel).Monomasti-
gote, uninucleate, with finger-like pseudopodia; flagellum long and
connected with nucleus; fresh water, soil or endocommensal.
Fig.
abed
115.
X1650 (Bishop)
(?); c,
Histomonas meleagridis as seen in smears from culture.
a, an amoeboid form; b, rounded form with axostyle
nucleus preparing to divide; d, a stage in nuclear division.
quail (Tyzzer).
This organism is the cause of enterohepatitis known as "black-
head, "an infectious disease, in young turkeys in which it is often
fatal and Smith (1895) discovered the organism
also in other fowls.
and considered it an amoeba. It invades and destroys the mucosa of
the intestine and caeca as well as the liver tissues. Trophozoites
voided in faeces by infected birds may become the source of new
infection when taken in by young birds with drink or food. Tyzzer
however demonstrated that the organism is transmissible from bird
to bird in the eggs of the nematode Heterakis gallinae. Bishop (1938)
made a cultural study of the organism. Dobell (1940) points out the
similaritybetween this flagellate and Dientamoeha fragilis (p. 368).
Wenrich (1943) made a comparative study of forms found in the
caecal smears of wild ring-neck pheasants and of chicks. The organ-
isms measured 5-SOfj. in diameter and possessed 1-4 flagella, though
often there were no flagella.
Genus Rhizomastix Alexeieff. Body amoeboid; nucleus central:
blepharoplast located between nucleus and posterior end; a long
ZOOMASTIGINA, RHIZOMASTIGINA 267
fiber runs from it to anterior end and continues into the flagellum;
without contractile vacuole; division in spherical cyst.
R. gracilis A. (Fig. 114, /). 8-14)u long; flagellum 20/i long; in
intestine of axolotles and tipulid larvae.
References
Becker, E. R. 1925 The morphology of Mastigina hylae (Frenzel)
from the intestine of the tadpole. Jour. Paras., Vol. 11.
Bishop, Ann 1938 Histomonas meleagridis in domestic fowls (Gal-
lus gallus). Cultivation and experimental infection. Parasitol-
ogy, Vol. 30.
DoBELL, C. 1940 Research on the intestinal Protozoa of monkeys
and man. X. Ibid., Vol. 32.
Lemmermann, E. 1914 Pantostomatinae. Siisswasserflora Deutsch-
H.L.
lands, etc.
Tyzzer, E. E. 1919 Developmental phases of the protozoon of
"blackhead" in Turkeys. Jour. Med. Res., Vol. 40.
Wenrich, D. H. 1943 Observations on the morphology of His-
tomonas from pheasants and chickens. Jour. Morph., Vol. 72.
Chapter 14
THEa protomonads
heterogeneous
of
possess one or two
Protozoa, mostly
lot of
and are composed
flagella
whose af-
parasitic,
finities to one another are very incompletely known. The body is in
268
PROTOMONADINA 269
In vertebrate In vertebrate
In invertebrate host
host host
Leptomonas and
Phytomonas (in plant)
Leishmania
Crithidia
Herpetomonas
Trypanosoma
and confined to, central Africa within a zone on both sides of the
equator where the vectors, Glossma palpalis and G. tachinoides (on
the west coastal region) live. Many wild animals have been found
naturally infected by the organisms and are considered to be reser-
voir hosts.
The chief lesions of infection are in the lymphatic glands and in
the central nervous system. In all cases, an extensive small-
there is
Q jO^O^
%
\ ^
j
^•-'
%®^ -?^o^' ^O
Fig. 122. Trypanosoma rhodesiense in a stained blood film of an inoculated
rat, X1150 (Kudo).
T. cruzi Chagas (Schizotrypanum cruzi C; T. triatomae Kofoid
and McCulloch) (Fig. 123). A small curved (C or U) form about
20/i long; nucleus central; blepharoplast conspicuously large, located
close to sharply pointed non-flagellate end; multiplication takes
r
I
place in the cells of nearly every organ of the host body; upon enter-
ing a host cell, the trypanosome loses its flagellum and undulating
membrane, and assumes a leishmania form which measures 2 to 5/^
in diameter; this form undergoes repeated binary fission, and a large
number of daughter individuals are produced; they develop sooner or
PROTOMONADINA 277
278 PROTOZOOLOGY
camels, cattle, swine, dogs, etc., which terminates in the death of
the host animal in from two weeks to a few months; wild animals
are equally susceptible; the disease occurs, of course, only in the
region in Africa where the tsetse flies live.
Watson. Similar to T.
T. peromysci lewisi; in Canadian deer mice,
Peromyscus maniculatus and others.
T. nahiasi Railliet. Similar to T. lewisi; in rabbits, Lepus do-
mesticus and L. cuniculus.
T. paddae Laveran and Mesnil. In Java sparrow, Munia oryzi-
vora.
IS
284 PROTOZOOLOGY
longitudinal division. Dogs are naturally infected with L. donovani
and may be looked upon as reservoir host. Vectors are Phlehotomus
argentipes and other species of Phlebotomus.
L. tropica (Wright). This is the causative organism of the Oriental
sore or cutaneous leishmaniasis. It has been reported from Africa
(mainly regions bordering the Mediterranean Sea), Europe (Spain
Italy, France, and Greece), Asia (Syria, Palestine, Armenia, South-
ern Russia, Iraq, Iran, Arabia, Turkestan, India, Indo-China, and
China), and Australia (northern Queensland). The organisms are
present in the endothelial cells in and around the cutaneous lesions,
located on hands, feet, legs, face, etc.
L. tropica is morphologically indistinguishable from L. donovani,
but some believe that it shows a wider range of form and size than
the latter. In addition to rounded or ovoid forms, elongate forms are
often found, and even leptomonad forms have been reported from
the scrapings of lesions. The insect vectors are Phlehotomus papa-
tasii (p. 282), P. sergenti and others. Direct transmission through
wounds in the skin also takes place. The lesion appears first as a
small papula on skin; it increases in size and later becomes ulcerated.
Microscopically an infiltration of corium and its papillae by lympho-
cytes and macrophages is noticed; in ulcerated lesions leishmania
bodies are found in the peripheral zone and below the floor of the
ulcers.
L. brasiliensis Vianna. This organism causes Espundia, Bubos, or
South American or naso-oral leishmaniasis, which appears to be
confined to South and Central America. It has been reported from
Brazil, Peru, Paraguay, Argentina, Uruguay, Bolivia, Venezuela,
Ecuador, Colombia, Panama, Costa Rica, and Mexico.
Its morphological characteristics are identical with those of L.
tropica, and a number of investigators combine the two species into
one. However, L. brasiliensis produces lesions mainly in the mucous
membrane of the nose, mouth and pharynx. Vectors appear to be
Phlehotomus intermedius and other species of the genus. Direct
transmission through wounds is also probable.
85m high.
Genus Rhipidodendron Stein. Similar to Cladomonas, but tubes
are fused lengthwise fresh water. ;
D. socialis K. (Fig. 129, g). Oval flagellum about 2-3 times the
body length; lorica yellowish or pale brown; broadly spindle in form;
about ISjLt long; pond water.
W_D. virgaria (Weisse) (Fig. 130, h). About 8m long; colony 200ju
high; pond water.
Genus Cephalothamnium Stein. Colonial; without but in-
lorica,
dividuals clustered at the end of a stalk which is colorless and rigid;
fresh water.
C. cyclopum S. (Fig. 130, i). Ovoid; 5-10/i long; attached to body
of Cyclops and also among plankton.
Genus Anthophysis Bory (Anthophysa). Colonial forms, some-
PROTOMONADINA 289
r ^ % ^
4 • •
8
Fig. 132. Retortanionas intestinalis, X1150. 1-3, living organisms;
4, 5, stained trophozoites; 6, a fresh cj^st; 7, 8, stained cysts (1-4,
Wenyon and O'Connor; 5, Dobell and O'Connor; 7, Jepps; 6, 8, Kudo).
Suborder I Monomonadina
Without axial organella
With 3 flagella Family 1 Trimastigidae (p. 294)
With 4 flagella
None undulates on body surface
Without cell-organ of attachment. Family 2 Tetramitidae (p.
. 296)
With rostellum Family 3 Streblomastigidae (p. 298)
One undulates on body surface Family 4 Chilomastigidae (p.
. . 298)
With more than 4 flagella Family 5 Callimastigidae (p. 299)
With axial organella
Without undulating membrane
Without cresta
Flagella not adhering to body
Without rostellum Family 6 Polymastigidae (p. 299)
293
294 PROTOZOOLOGY
With rostellum Family 7 Oxymonadidae (p. 301)
Flagellar cords on body surface
Family 8 Dinenymphidae (p. 302)
With cresta Family 9 Devescovinidae (p. 303)
With undulating membrane. . . .Family 10 Trichomonadidae (p. 308)
o-
POLYMASTIGINA 297
P \f' % %
% '^- "0 •
2 7 8 9 10
Fig. 138. a, Streblomaslix strix, X1030; b, its anterior end, showing the
rostellum, blepharoplast, sucking cup and flagella (Kidder); c, Calli-
masiix frontalis, X1500 (Braune); d, C. equi, XUOO (Hsiung); e, Poly-
mastix melolonthae, X546 (Hamburger); f, Eutrichomastix serpentis,
X1450 (Kofoid and Swezy); g, E. hatracJiorum, X 1350 (Dobell); h, E.
axostylis, X2000 (Kirby); i, Hexamastix termopsidis, X2670 (Kirby);
j, H. batrachorum, XlOOO (Alexeieff); k, Protrichomonas legeri, XlOOO
(Alexeieff); 1, Monocercomonas bufonis, X1670 (Alexeieff).
papilla
ant. flagella
ant. lamella
bleph. group
nucL rhiz.
parab. f il.
parab. body
cresta
chrom. mass
nucl. memb.
chr. cone in ax.
parab. spiral
chromoph. element
of pb.
axostyle
tr. flagellum
\( aI
r 1 t
Fig. 144. TricJiomonas hominis, X1150 (Kudo). 1, living and
2, 3, stained trophozoites.
Suborder 2 Diplomonadina
T. rotans Klebs (Fig. 147, g). Broadly oval; posterior half highly
flattened; 2 long and 2 short on each of 2 cytostomes; stag-
flagella
nant water.
Genus Gyromonas Seligo. Free-swimming; small; form constant,
flattened; slightly spirally coiled; 4 flagella at anterior end; cyto-
stome not observed; fresh water.
G. ambulans S. (Fig. 147, h). Rounded; 8-1 5/i long; standing
water.
Genus Trigonomonas Klebs. Free-swimming; pyriform, plastic;
cytostome on either side, from anterior margin of which arise 3
flagella; flagella 6 in all; 2 nuclei situated near anterior end; move-
ment rotation; holozoic; fresh water.
POLYMASTIGINA 315
Suborder 3 Polymonadina
References
Cleveland, 1934 The wood-feeding roach, Cryptocer-
L. R. et al.
cus, its Protozoa, and the symbiosis between Protozoa and
roach. Mem. Amer. Acad. Arts and Sci., Vol. 17.
Davis, H. S. 1943 A new polymastigine flagellate, Costia pyri-
formis, parasitic on trout. Jour. Parasit., Vol. 29.
DoBELL, C. 1934 Researches on the intestinal Protozoa of monkeys
and man. VI. Parasitology, Vol. 26.
POLYMASTIGINA 317
—
— 1935 VII. Parasitology, Vol. 27.
1939 The common flagellate of the human mouth, Tri-
chomonas tenax (O.F.M.): its discovery and its nomenclature.
Ibid., Vol. 31.
and F. W. O'Connor 1921 The intestinal Protozoa of man.
London.
Feo, L. G. 1944 The incidence and significance of Trichomonas
vaginalis in the male. Amer. Jour. Trop. Med., Vol. 24.
Grasse, p. p. 1926 Contribution a I'^tude des Flagelles parasites.
Arch. zool. exp. g^n.. Vol. 65.
HoGUE, Mary J. 1943 The effect of Trichomonas vaginalis on tissue-
culture cells. Amer. Jour. Hyg., Vol. 37.
KiRBY, H. 1930, 1931 Trichomonad flagellates from termites. I, II.
Uni. Cal. Pub. Zool., Vols. 33, 36.
1941, 1942 Devescovinid flagellates of termites. I, II, III.
Ibid., Vol. 45.
1943 Observations on a trichomonad from the intestine of
man. Jour. Parasit., Vol. 29.
KoFOiD, C. A. and Olive Swezy 1915 Mitosis and multiple fission
in trichomonad flagellates. Proc. Amer. Acad. Arts and Sci.,
Vol. 51.
1920 On the morphology and mitosis of Chilomas-
tix mesnili (Wenyon), a common flagellate of the human intes-
tine. Uni. Cal. Pub. Zool., Vol. 20.
1922 Mitosis and fission in the active and encysted
phases of Giardia enterica etc. Ibid., Vol. 20.
McNeil, E., W. R. Hinshaw and C. A. Kofoid 1941 Hexamita
meleagridis sp. nov. from the turkey. Amer. Jour. Hyg., Vol. 34.
Rees, C. W. 1938 Observations on bovine venereal trichomoniasis.
Veterin. Med., Vol. 33.
Stabler, R. M., L. G. Feo and A. E. Rakoff 1941 Implantation
of intestinal trichomonads {T. hominis) into the human vagina.
Amer. Jour. Hyg., Vol. 34.
Sutherland, J. L. 1933 Protozoa from Australian termites. Quart.
Jour. Micr. Sci., Vol. 76.
Wenrich, D. H. 1944 Comparative morphology of the trichomo-
nad flagellates of man. Amer. Jour. Trop. Med., Vol. 24.
1944a Morphology of the intestinal trichomonad flagellates
in man and of similar forms in monkeys, cats, dogs, and rats.
Jour. Morph., Vol. 74.
Wenyon, C. M. 1926 Protozoology, Vol. 1. London.
Chapter 16
ALL members of
canal termites,
of
this order are inhabitants of the
cockroaches, and woodroaches.
aUmentary
The cyto-
plasmic organization is of high complexity, although there is only
a single nucleus. Flagella are numerous and have their origin in the
blepharoplasts located in the anterior region of body. In many spe-
cies, there exists a true symbiotic relationship between the host
termite and the protozoans (p. 25). Method of nutrition is either
holozoic or saprozoic (parasitic). Bits of wood, starch grains, and
other food material are taken in by means of pseudopodia.
Asexual reproduction is by longitudinal fission; multiple division
has also been noted in some species under certain conditions, while
sexual reproduction has not been observed. Encystment occurs in
some genera of Lophomonadidae and certain species inhabiting
woodroaches, in which moulting of the host insect leads to encyst-
ment. Because of the peculiarity and complexity of their structures
and also of their common occurrence in termites, the Hypermastigina
have been frequently studied.
Body without segmented appearance
Flagella in spiral rows Family 1 Holomastigotidae
Flagella not arranged in spiral rows
Flagella in one or more anterior tufts
1 tuft of flagella Family 2 Lophomonadidae (p. 320)
2 tufts of flagella Family 3 Hoplonymphidae (p. 322)
4 tufts of flagella Family 4 Staurojoeninidae (p. 324)
Several tufts (loriculae) Family 5 Kofoidiidae (p. 324)
Flagella not arranged in tufts
Posterior part without flagella
Family 6 Trichonymphidae (p. 324)
Flagella over entire body.. .Family 7 Eucomonymphidae (p. 326)
Body with segmented appearance. .Family 8 Teratonymphidae (p. 326)
gut.
References
Cleveland, L. R. 1925 The
effects of oxygenation and starvation
on the symbiosis between the termite, Termopsis, and its intes-
tinal flagellates. Biol. Bull., Vol. 48.
and others 1934 The wood-feeding roach, Cryptocercus, its
Protozoa, and the symbiosis between Protozoa and roach. Mem.
Amer. Acad. Arts and Sci., Vol. 17.
HYPERMASTIGINA 327
THE
and,
members of this class do not possess
therefore, are capable of forming pseudo podia
any definite pellicle
(p. 41).
The term 'amoeboid' is often used to describe their appearance.
The pseudopodia serve ordinarily for both locomotion and food-
capturing. The peripheral portion of the body shows no structural
differentiation in Amoebina, Proteomyxa, and Mycetozoa. Internal
and external skeletal structures are variously developed in other
orders. Thus, in Testacea and Foraminifera, there is a well-devel-
oped test or shell that usually has an aperture, through which the
pseudopodia are extruded; in Heliozoa and Radiolaria, skeletons of
various forms and materials are developed.
The cytoplasm is, as a rule, differentiated into the ectoplasm and
the endoplasm, but this differentiation is not constant. In Radio-
laria, therea perforated membranous central capsule which marks
is
the border line between the two cytoplasmic regions. The endoplasm
contains the nuclei, food vacuoles, various granules, and contractile
vacuoles. The majority of Sarcodina are uninucleate, butnumerous
species of Foraminifera and Mycetozoa are multinucleate in certain
phases during their development. In the family Paramoebidae, there
occurs a peculiar secondary nucleus.
The Sarcodina are typically holozoic. Their food organisms are
Protozoa, small Metazoa and Protophyta, which present themselves
conspicuously in the cytoplasm. One or more contractile vacuoles
are invariably present in forms inhabiting the fresh water, but absent
in parasitic forms or in those which live in the salt water.
328
SARCODINA, PROTEOMYXA 329
yhnK'
334 PROTOZOOLOGY
and anastomosing, colorless; nucleus inconspicuous; one or more
contractile vacuoles, small fresh water.
;
References
Cash, J. 1905 The British freshwater Rhizopoda and Heliozoa. Vol.
1. London.
DoBELL, C. 1913 Observations on the life-history of Cienkowski's
Arachnula. Arch. f. Protistenk., Vol. 31.
DoFLEiN, F. and E. Reichnow 1929 Lehrbuch der Protozoenkunde.
Jena.
DuBOSCQ, 0. 1921 Labyrinthomyxa sauvageaui n. g., n. sp., pro-
teomyxee parasite de Laminaria lejolisii Sauvageau. C. r. soc.
biol., Paris. Vol. 84.
KtJHN, A. 1926 Morphologie der Tiere in Bildern. H. 2; T. 2.
Rhizopoden.
Leidy, J. 1879 Freshwater Rhiozopods of North America. Report
U. S. Geol. Survey. Vol. 12.
RosKiN, G. 1927 Zur Kenntnis der Gattung Pseudospora Cienkow-
ski. Arch. f. Protistenk., Vol. 59.
Valkanov, a. 1940 Die Heliozoen und Proteomyxien. Ibid., Vol.
93.
Young, E. L. 1943 Studies on Labyrinthula, the etiologic agent of
the wasting disease of eel-grass. Amer. Jour. Bot., Vol. 30.
ZoPF, W. 1887 Handhuch der Botanik (A. Schenk). Vol. 3.
Chapter 18
THE Mycetozoaknown
fungi,
had been considered
being,
be
Mj^xomycetes, or
as
to
Myxogasteres, the
closely related to the
335
336 PROTOZOOLOGY
When lack of food material occurs, the Plasmodium undergoes
changes and develops sporangia. The first indication of this process
is the appearance of lobular masses of protoplasm in various parts
of the body (/, g). These masses are at first connected with the stream-
ing protoplasmic thickenings, but later become completely segre-
gated into young sporangia. During the course of sporangium-for-
mation, foreign bodies are thrown out of the body, and around each
the base, hypothallus, for the stalk. With these changes the interior
of the sporangium becomes penetrated by an anastomosing network,
capillitium, of flat bands which are continuous with the outer cover-
ing (^). Soon after the differentiation of these protective and sup-
porting structures, the nuclei divide simultaneously by mitosis and
the cytoplasm breaks up into many small bodies. These uninucleate
bodies are the spores which measure 3-20ju in diameter and which
soon become covered by a more or less thick cellulose membrane (j),
variously colored in different species.
The mature sporangium breaks open sooner or later and the
spores are carried, and scattered,by the wind. When a spore falls
in water, its membrane ruptures, and the protoplasmic contents
emerge as an amoebula {k, I). The amoebula possesses a single vesic-
idar nucleus and contractile vacuoles, and undergoes a typical amoe-
boid movement. It presently assumes an elongate form and pro-
trudes a flagellum from the nucelated end, thus developing into a
myxofiagellate (zoospore or swarmer) (m, n) which undergoes a pe-
culiar dancingmovement and is able to form short, pointed pseudo-
podia from the posterior end. It feeds on bacteria, grows and multi-
plies by binary fission (o-q). After a series of division, the myxo-
fiagellate may encyst and becomes a microcyst (r). Wlien the micro-
cyst germinates, the content develops into a myxamoeba
(s) which,
Family 8 Reticulariidae
Phytomyxinea Poche
These organisms which possess a large multinucleate amoeboid
body, are parasitic in various plants and also in a few animals.
^4}^
References
DE Bary 1864 Die Mycetozoa. Leipzig.
Hagelstein, R. 1944 The Mycetoza of North America. New York.
Jahn, E. 1901-1920 Myxomycetenstudien. I to X. Ber. Deutsch.
Bot. Ges., Vols. 19, 20, 22-26, 29, 36 and 37.
Jones, P. M. 1928 Morphology and cultural study of Plasmodio-
phora brassicae. Arch. f. Protistenk., Vol. 62.
Karling, J. S. 1942 The Plasmodiophorales. New York.
Lister, A. 1925 A monograph on the Mycetozoa. 3rd ed. London.
MacBride, T. H. 1922 North American slime molds. 2nd ed. New
York.
and G. H. Martin 1934 The Myxomycetes. New York.
Chapter 19
THEnoAmoebinaorshow asurrounding
is pellicle
very
test,
little cortical differentiation.
343
344 PROTOZOOLOGY
Family 1 Dimastigamoebidae Wenyon
The members of the two genera placed both
in this family possess
amoeboid and flagellate phases {diphasic).
In the former, the organ-
ism undergoes amoeboid movement by means of lobopodia and in
the latter the body is more or less elongated. Binary fission seems to
take place during the amoeboid phase only. Thus these are diphasic
protozoans, in which the amoeboid stage predominates over the
flagellate. The amoeboid phase resembles a 'limax' amoeba; under
natural circumstances, it is often exceedingly difficult by observing
the amoeboid stage only, to determine whether they belong to this
family or the family Amoebidae.
Fig. 160. a-c, trophozoite, flagellate phase and cj^st (all stained) of
Dimastig amoeba gruberi, x750 (Alexeieff); d-f, similar stages of D.
bistadialis, X750 (Kiihn); g-j, trophozoite, flagellate phase, cyst and
excystation of Trimastigamoeba philippi7iensis X950 (Whitemore).
Ms.
350 PROTOZOOLOGY
on various Protozoa and also small invertebrates; may-
oles; feeds
take 20 Paramecium within one food-cup; plasmotomy into 2-6
smaller individuals. North America (North Carolina (Wilson); Vir-
ginia(Kepner and Edwards) Tennessee and New Jersey (Schaeff er)
;
«)
tion rate varied from 0.2 to 53 per cent, averaging 11.6 per cent,
which justifies Craig's (1926) earlier estimate that about 10 per cent
of the general population harbor this protozoan. An acute infection
by E. histolytica is accompanied by dysenterj^, while in chronic cases
or in convalescence, the host may void infectious cysts without
suffering from the infection himself. Such a person is known as a
cyst-carrier or -passer.
The trophozoite if voided in faeces perish in a comparatively short
time. The dissemination of infection is exclusively carried on by the
cyst. Viable cysts may be transmitted (1) by contamination of food
through contact with contaminated water or through unsanitary
habit of food handlers who are cyst-carriers; (2) by droppings of
flies and cockroaches which, as noted below, contain viable cysts for
358 PROTOZOOLOGY
unstained cysts might be dead or uninfectious is unknown. But as
Wenyon and O'Connor wrote, "if we accept the eosin test as a
criterion and regard all unstained cysts as living, the error in judg-
ment willbe on the safe side." Root found neutral red in 1:10,000
dilution to give a slightly larger proportion of stained cysts than
eosin. Frye and Meleney's (1936) comparative study leads one to
look upon this method as a fairly dependable one.
(b) Cultivation test. Improved cultural technique now brings
about easily excystment of viable cysts in a proper culture medium.
For example, Yorke and Adams (1926) obtained in 24 hours "a
plentiful growth of vegetative forms" from cysts in Locke-egg-serum
medium (p. 717). Snyder and Meleney (1941) note recently that the
excystation does not take place in various culture media unless liv-
ing bacteria were added or oxygen concentration of the media was
decreased. Animal infection method has not been used much, as
experimental animals (cats) show individual difference in suscepti-
bility. Some of the published results are summarized below. The
testing method used is indicated by: a for eosin test or h for cultiva-
tion test and is given after the name of the investigators.
1. Cysts in faeces kept in a covered container. All cysts disap-
all dead on the 13th day (Kuenen and Swellengrebel; a). Viable for
for 153 days (Boeck; a). Alive for 10 and 17 days at 16-20° and 0°C.
respectively (Yorke and Adams; 6); for 3. 10, 30, and 90 days at
30°, 20°, 10° and 0°C. respectively (Chang and Fair; b).
3. Cysts in relation to high temperatures. Cysts are killed at
1.
AMOEBINA 561
-0,
X
,©
m.'
:^
.o^
intestine of guinea-pigs.
E. muris (Grassi). Similar to E. coli; in intestine of rats and mice.
E. gallinarum Tyzzer. In fowl's intestine; cysts octo nucleate.
AMOEBINA 365
w
§> « jrf'
''-^
© m u:
Fig. 171. lodamoeba butschlii, X1150 (Kudo). 1, a living amoeba; 2-5,
©
stained trophozoites; 4, 5, somewhat degenerating trophozoites; 6, a fresh
cyst; 7-10, stained cysts.
Several species.
E. nana (Wenyon and O'Connor) (Fig. 172). The trophozoite
measures 6-15jli in diameter; fairly active monopodial movement by
forming a broad pseudo podium; when stationary pseudo podia are
formed at different points; endoplasm is granulated and contains
bacteria as food particles; the vesicular nucleus, 1.5-3)u in diameter,
iscomposed of a delicate membrane with a few chromatin granules,
and a large irregularly shaped endosome.
considered as a commensal.
E. gregariniformis (Tyzzer). In caecum of fowls; 4-12)u in diam-
eter; cysts uninucleate.
E. ranarum Epstein and Ilovaisky. In colon of frogs; cyst octo nu-
cleate, up to 25ju in diameter.
E. blattae Lucas. In colon of cockroaches; 3-15^ long; cyst with
more than one nucleus.
Genus Dientamoeba Jepps and Dobell. Small amoeba; number of
binucleate trophozoites often greater than that of uninucleate
forms; nuclear membrane delicate; endosome consists of several
chromatin granules embedded in plasmosomic substances and
connected with the membrane by delicate strands (Fig, 164, e); in
colon of man.
368 PROTOZOOLOGY
D. fragilis J. and D. (Fig. 173). The trophczoite is amoe-
actively
boid; i-18fx (average 5-12/i) in diameter; progressive movement;
cj^toplasm well differentiated; endoplasm granulated contains bac-
teria in food vacuoles; nucleus onty faintly visible; 1 or 2 nuclei, the
ratio is variable; in some material binucleate forms may be 80% or
more, while in others uninucleate forms may
predominate; nucleus
is made up of a delicate membrane and a large endosome (more than
12 3 4
Fig. 174. a-f, Dobellina viesnili (Bishop and Tate): a, b, uni- and multi-
nucleated stained amoebae, X2200; c, a trinucleate (stained) amoeba in
which the nuclei are dividing, X1760; d-f, stained cysts with 2, 4, and 6
nuclei, X1760 (Bishop and Tate); g, h, stained trophozoite and cyst of
Schizamoeba salmonis, X1070 (Davis); i, j, Hydramoeha hydroxena (Rey-
nolds and Looper): i, a heavily infected Hydra oligadis, X95; j, part of a
section of an infected host hydra, showing a trophozoite feeding on ecto-
dermal cells, X470; k, a stained individual of Paramoeba -pigmentifera,
with its nucleus in the center, X800 (Janicki).
References
the body can completely be withdrawn. The shell has usually a single
aperture through which pseudopodia protrude, and varies in shape
and structure, although a chitinous or pseudochitinous membrane
forms the basis of all. It may be thickened, as in Arcella and others,
or composed of foreign bodies cemented together as in Difflugia,
while in Euglypha siliceous platelets or scales are formed in the
endoplasm and deposited in the shell.
The cytoplasm is ordinarily differentiated into the ectoplasm and
endoplasm. The ectoplasm is conspicuously observable at the aper-
ture of the shell where filopodia or slender ectoplasmic lobopodia
are produced. The endoplasm is granulated or vacuolated and con-
tains food vacuoles, contractile vacuoles and nuclei. In some forms
there are present regularly in the cytoplasm numerous basophilic
granules which are known as 'chromidia' (p. 37).
Asexual reproduction is either by longitudinal fission in the forms
with soft tests, or by transverse division or budding, while in others
multiple division occurs. Encystment is common. Sexual reproduc-
tion b}^ amoeboid or flagellate gametes has been reported in some
species.
The testaceans are mostly inhabitants of fresh water, but some
live in saltwater and others are semi-terrestrial, being found in
moss or moist soil, especiallj^ peaty soil.
and membranous
Shell simple
some anastomosing
Filopodia, in Family 1 Gromiidae
Pseudopodia filose, simplj^ branched Family 2 Arcellidae (p. 378)
Shell with foreign bodies, platelets, or scales
With foreign bodies Family 3 Difflugiidae (p. 384)
With platelets or scales Family 4 Euglyphidae (p. 389)
374
TESTACEA 375
branching and anastomosing; cytoplasm with numerous
motile
granules; fresh or salt water. Man}^ species.
mm
Fig. 175. a, Gromia jiuvialis, X120 (Dujardin); b, G. ovoidea, X50
(Schultze); c, G. nigricans, x200 (Cash and Wailes); d, Microgromia
sociahs, X170 (Cash); e, Microcometes paludosa, X670? (Penard)-
f, Artodiscus saltans, X670 (Penard); Schultzella
g, diffluens, Xl2()
(Rhumbler).
376 PROTOZOOLOGY
G.fluvialis D. (Fig. 175, a). Test spherical to subspherical smooth;
tation.
Genus Cryptodifflugia Penard, Small test yellowish to brownish;
Difflugia-like in general appearance, compressed; with or without
foreign bodies; pseudopodia long, acutely pointed; fresh water.
C. oviformis P. (Fig. 178,/). Test ovoid; without foreign bodies;
crown hemispherical; aperture truncate; cytoplasm with chloro-
phyllous food particles; 16-20iu by 12-15/1 in marshy soil.
;
etc.
D. oblonga Ehrenberg (D. pyriformis Perty) (Fig. 180, a). Test
pyriform, flask-shaped, or ovoid; neck variable in length; fundus
rounded, with occasionally 1-3 conical processes; aperture terminal,
;
TESTACEA 385
Several varieties.
D. urceolata Carter ^Fig. 180, b). A large ovoid, rotund test, with
a short neck and a rim around aperture; 200-230/i by 1 50-200/1
THE
most
Foraminifera are comparatively large Protozoa, living al-
They were very abundant in geo-
exclusively in the sea.
logic times and the forms are important in applied geology
fossil
(p. 10). The majority live on ocean bottom, moving about slug-
gishly over the mud and ooze by means of their pseudo podia. Some
are attached to various objects on the ocean floor, while others are
pelagic.
The cytoplasm is ordinarily not differentiated into the two zones
and streams out through the apertures, and in perforated forms
through the numerous pores, of the shell, forming rhizopodia which
are fine and often very long and which anastomose with one another
to present a characteristic appearance (Fig. 5). The streaming move-
ment of the cytoplasm in the pseudopodia are quite striking; the
granules move toward the end of a pseudopodium and stream back
along its periphery. The body cytoplasm is often loaded with brown
granules which are apparently waste matter and in some forms such
as Peneroplis pertusus these masses are extruded from the body
from time to time, especially prior to the formation of a new cham-
ber. Contractile vacuoles are usually not found in the Foraminifera.
The test of the Foraminifera varies greatly inform and structure.
It may show various colorations —
orange, red, brown, etc. The ma-
jority measure less than one millimeter, although larger forms may
frequently reach several millimeters. The test may be siliceous or
calcareous and in some forms, various foreign materials, such as
sand-grains, sponge-spicules, etc. which are more or less abundantly
found where these organisms live, are loosely or compactly cemented
together by pseudochitinous or gelatinous substances. Certain forms
show a specific tendency in the selection of foreign materials for the
test (p. 40). Siliceous tests are comparatively rare, being found
in some species of Miliolidae inhabiting either the brackish water or
deep sea. Calcareous tests are sometimes imperforated, but even in
such cases those of the young are always perforated. By far the ma-
jority of the Foraminifera possess perforated calcareous tests. The
thickness of the shell varies considerably, as do also the size and
number of apertures, among different species. Frequently the per-
forations are very small in the young and later become large and
coarse, while in others the reverse may be the case.
394
FORAMINIFERA 395
The form of the shell varies greatly. In some there is only one
chamber composed of a central body and radiating arms which repre-
sent the material collected around the pseudopodia, as in Rhabdam-
mina (Fig. 185, a) or of a tubular body alone, as in Hyperammina (Fig.
,
A B r
{2). They grow into mature megalo spheric forms which measure
60-72)U in diameter. Two to four such individuals become associated
and transform into "fertilization cyst." (S). The nucleus in each
individual divides twice or occasionally three times and thus formed
multinucleate bodies escape from the tests within the cyst envelope
where many gametocytes are produced by multiple fissions. Each
gametocyte which contains 12 chromosomes divides into two amoe-
boid haploid gametes by meiosis. Gametes developed from different
parents presumably undergo fusion in pairs and zygotes are pro-
Family 1 Astrorhizidae
Family 15 Trochamminidae
Bj% a ^"
d ...;;:
k J 1
# @
{Polysiomella Lamarck)
FORAMINIFERA 403
Family 30 Buliminidae
Family 35 Halkyardiidae
References
Brady, B. H. 1884 Report on the Foraminifera dredged by H.M.S.
Challenger, during the years 1873 to 1876. Rep. Voy. Challenger,
Vol. 9.
CusHMAN, J. A. 1940 Foraminifera: their classification and economic
use.Third edition. Cambridge, Mass.
Myers, E. H. 1935 The life history of Patellina corrugata William-
son, a foraminifer. Bull. Scripps Inst. Oceanogr. Uni. Calif.,
Tech. Ser., Vol. 3.
1936 The life-cycle of Spirillina vivipara Ehrenberg, with
notes on morphogenesis, systematics and distribution of the
Foraminifera. Jour. Roy. Micr. Soc, Vol. 56.
1938 The present state of our knowledge concerning the life
cycle of the Foraminifera. Proc. Nat. Acad. Sci., Vol. 24.
1940 Observations on the origin and fate of flagellated
gametes in multiple tests of Discorbis (Foraminifera). Jour. Ma-
rine Biol. Ass. United Kingdom, Vol. 24.
Rhumbler, L. 1904 Systematische Zusammenstellung der rezenten
Reticulosa (Nuda u. Foraminifera). Arch. f. Protistenk., Vol. 3.
Chapter 22
Subclass 2 Actinopoda Calkins
THE Actinopoda
Without
are divided into
central capsule
two orders as follows:
Order 1 Heliozoa
With central capsule Order 2 Radiolaria (p. 417)
406
ACTINOPODA, HELIOZOA 407
tion studied by Belaf (p. 164); Looper (1928) studied its food re-
actions.
A. vesiculata Penard. Ectoplasm with saccate secondary vesicles,
extending out of body surface between axo podia; nucleus central,
with many endosomes; 25-30/1 in average diameter; fresh water.
Genus Actinosphaerium Stein. Spherical; ectoplasm consists al-
most entirely of large vacuoles in one or several layers; endoplasm
with numerous small vacuoles; numerous nuclei; axial filaments end
in the inner zone of ectoplasm. 2 species.
A. eichhorni Ehrenberg (Figs. 6; 190, c). Numerous nuclei scattered
in the periphery of endoplasm; 2 or more contractile vacuoles, large;
axial filaments arise from a narrow zone of dense cytoplasm at the
border line between endoplasm and ectoplasm; body large, diameter
200-300/i, sometimes up to 1 mm.; nuclei 12-20/z in diameter; among
vegetation in freshwater bodies.
A. arachnoideum Penard. Ectoplasm irregularly vacuolated; no
distinct endoplasmic differentiation; nuclei smaller in number; pseu-
dopodia of 2 kinds; one straight, very long and the other filiform,
and anastomosing; 70-80/x in diameter; fresh water.
Genus Camptonema Schaudinn. Spheroidal; axial filaments of
axopodia end in nuclei about 50 in number; vacuoles numerous and
small in size; salt water.
About ISOyu in diameter.
C. nutans S. (Fig. 190, d).
Genus Oxnerella Dobell. Spherical; cytoplasm indistinctly dif-
ferentiated; eccentric nucleus with a large endosome; axial filaments
take their origin in the central granule; no contractile vacuole;
nuclear division typical mitosis (Fig. 58).
0. maritima D. (Fig. 58). Small, 10-22/i in diameter; solitary,
floating or creeping; salt water.
II. Arch.
Protistenk., Vols. 46, 48.
f.
THE Radiolaria
ocean known
floor is
are pelagic in various oceans. A vast area of the
be covered with the ooze made up chiefly
to
of radiolarian skeletons. They seem to have been equally abundant
during former geologic ages, since rocks composed of their skeletons
occur in various geological formations. Thus this group is the second
group of Protozoa important to geologists.
The body is generally spherical, although radially or bilaterally
symmetrical forms are also encountered. The cytoplasm is divided
distinctly into two regions which are sharply delimited by a mem-
branous structure known as the central capsule. This is a single or
double perforated membrane of pseudochitinous or mucinoid nature.
Although its thickness varies a great deal, the capsule is ordinarily
very thin and only made visible after addition of reagents. Its shape
varies according to the form of the organism; thus in spherical forms
it is spherical, in discoidal or lenticular forms it is more or less ellips-
Family 1 Actineliidae
RADIOLARIA 421
Genus Sphaerocapsa
2 or 6 larger spines
2 enormously large conical sheathed spines
, Family 7 Diploconidae
Genus Diploconus
6 large spines Family 8 Hexalaspidae
Genus Thalassothamnus
A latticed skeleton, with branching and thorny spines
Family 5 Orosphaeridae
Genus Orosphaera
Solitary, skeleton complex, often concentric
Central capsule and skeleton spherical Family 6 Sphaeroidae
Genus Collosphaera
-^e^vii 'jm^d%'} w
"
- -. '«
©1
1 '
\
\
With skeleton
Without a complete latticed skeleton
Skeleton a basal tripod Family 2 Plectoidae
Genus Phormobothrys
Suborder 4 Triplylea Hertwig
Without skeleton; with isolated spicules
Skeleton consists of radial hollow rods and fine tangential needles
Family 1 Aulacanthidae
With skeleton
1-2 (concentric) usually spherical skeletons
Outer lattice skeleton with triangular or areolar meshes
Family 3 Sagosphaeridae
Genus Sagenoscene
One lattice skeleton with hollow radial bars
Family 4 Aulosphaeridae
Genus Cannosphaera
One skeleton, simple, but variable in shape; bilaterally symmetrical
Skeleton with fine diatomaceous graining . . Family 6 Challengeridae
RADIOLARIA 425
Skeleton similar to the last, but the base of each radial spine sur-
rounded by pores Family 9 Circoporidae
Chapter 24
Class 3 Sporozoa Leuckart
The spore which contains neither a polar capsule nor a polar fila-
the trophic life of the individual. In the forms which invade two host
427
:
428 PROTOZOOLOGY
animals to complete their development, there occur naked sporo-
zoites instead of spores.
The infection of a new host begins with the entrance of mature
spores through mouth, or with the introduction of the sporozoites
by blood-sucking invertebrates directly into the blood stream. The
sporozoites enter specific host cells and there grow at the expense of
the latter. In the Coccidia and the Haemosporidia, the trophozoite
continues its intracellular existence, but in the Gregarinida it leaves
the host and grows in an organ cavity. Except Eugregarinina,
cell
of gametes (d-f) which unite in pairs (g). The zygotes thus formed
develop into spores, each possessing 8 sporozoites (A). Meanwhile
the host pupa emerges as an adult mosquito, and the spores which
:
become set free in the lumen of the tubules pass into the intestine,
from which they are discharged into water. Larvae swallow the
spores and acquire infection.
Eugregarinina are divided into 2 tribes
Trophozoite not septate Tribe 1 Acephalina
Trophozoite septate Tribe 2 Cephalina (p. 440)
The acephalines are mainly found in the body cavity and organs
associated with it. The infection begins by the ingestion of mature
spores by a host, in the digestive tract of which the sporozoites are
set free and undergo development or make their way through the
gut wall and reach the coelom or various organs such as seminal
vesicles. Young trophozoites are intracellular, while more mature
forms are either intracellular or extracellular.
434 PROTOZOOLOGY
zooamylon bodies; cysts ovoid, 95/i by 84^1; spores 13.3/i by 5^.
R. porrecta Schmidt (Fig. 203, 6, c). In seminal vesicles of Lum-
hricus ruhellus and Helodrilus foetidus; extremely long with an en-
larged head; up to 2.5 mm. by 32-36m; sluggish; endoplasm granu-
lated, filled with oval {4n by 2-3/x) paraglycogen grains; nucleus 17-
25)u in diameter; spores 27.7-28/x by 12iu; sporozoites 13-18/x by 3-5/i.
100m long.
Genus Albertisella Cognetti. Mature trophozoites cup-shaped,
with anterior sucker with a smooth wall; nucleus at its bottom. One
species.
A. crater C. In seminal vesicles of Pheretima sermowaiana.
Sporadins solitary
Epimerite simple knob-like
Cysts with several ducts Family 6 Leidyanidae (p. 445)
Cysts without or with one duct
Family 7 Monoductidae (p. 445)
Epimerite not simple knob-like
Epimerite cup-shaped or digitate
Epimerite cup-shaped Family 8 Menosporidae (p.
. . . 448)
Epimerite digitate. ..Family 9 Dactylophoridae (p.
. 448)
Epimerite otherwise
Spore hat-shaped Family 10 Stylocephalidae (p. 450)
Spore of other shapes
Spore with spines. .Family 11 Acanthosporidae (p. 451)
Spore without spines
Family 12 Actinocephalidae (p. 452)
Two host species involved Family 13 Porosporidae (p. 455)
phrya desmaresii.
SPOROZOA, GREGARINIDA 445
Family 6 Leidyanidae
Similar to the last two families; but sporadins are solitary and
epimerite simple knob-like; cysts with several sporoducts.
Genus Leidyana Watson. Solitary; epimerite a simple globular
knob; cysts with ducts; spores dolioform.
sessile
L. erratica (Crawley) (Fig. 207,/). Sporadins up to 500m by 160m;
cysts about 350m in diameter; membrane about 30m thick; 1-12
sporoducts; spores extruded in chains, 6m by 3m; in gut of Gryllus
ahhreviatus and G. pennsylvanicus.
XSO (L6ger).
P. striatus Leger and Duboscq (Fig. 209, s). Sporadins 150^ long;
nucleus in protomerite; cysts spherical; in gut of larvae of Ptychop-
tera contaminata.
Genus Stylocystis Leger. Epimerite a sharply pointed, curved
process; spores biconical.
S. praecox L. (Fig. 209, t). Sporadins up to SOOyulong; cysts ovoidal,
200ju long; spores Sju by 5^ in gut of larval Tanypus sp.
Fig. 214. a-c, Selenidium potainillae (a, X420; b, cyst with spores,
X330; c, spore) (Mackinnon and Ray); d-f, Meroselenidium keilini
(d, sporadin, X670; e, f, different views of spore, X930) (Mackinnon
and Ray); g-j, Machadoella triatomae (g, a schizont, X1420; h, i, a single
and associated sporadins, X710; j, spore, X1920) (Reichenow); k, 1,
Syncystis ynirabilis: k, a cyst, X470 (Steopoe); 1, spore (Schneider);
m, Mattesia dispora, X1480 (Naville); n, Lipotropha macrospora, X800
(Keilin).
462 PROTOZOOLOGY
Genus Meroselenidium Mackinnon and Ray. Schizogony intra-
cellular, initiated by formation of small masses which give rise
to merozoites; about 20 spores from a pair of gametocytes; spores
with numerous sporozoites. One species.
M. keilini M. and R. (Fig. 214, d-f). Large schizonts about 150m
by 30/x; sporadins free in gut 200-300m by 40-70/i; paired gameto-
cytes 85/1 by 40m; spores 26-28m by 14- 16m, bivalve (?), trans-
verse ridges, with many sporozoites; in gut of Potomilla reniformis.
Genus Machadoella Reichenow. Nematode-like, rigid; simple
rounded anterior end; thick pellicle, longitudinally striated; schi-
zogony in vermiform stage; head to head association of gameto-
cytes; cysts with 3-6 spores, each with 8 sporozoites.
M. triatomae R. (Fig. 214, g-j). Schizonts about 55m long; game-
tocytes 100-120m long; schizogony into 6-8 merozoites; cysts with
3-6 spores; spore 10-1 1m by 7-7. 5m; in Malpighian tubules of
Triatoma dimidiata (of Guatemala)
References
DoFLEiN, F. and E. Reichenow 1929 Lehrbuch der Proiozoen-
kunde. 5th edition. Jena.
Labbe, a. 1899 Sporozoa. In Das Tierreich. Part 5.
Naville, a. 1931 Les Sporozoaires. Mem. d'hist. nat. de Geneve,
Vol. 41.
Wenyon, C. M. 1926 Protozoology. Vols. 1, 2.
THE vertebrates
Coccidia show a wide
all and higher invertebrates
zoological distribution,
alike.
attacking
The majority
are parasites of the epithelium of the digestive tract and its asso-
ciated glands. Asexual reproduction by schizogony and sexual
is
nuclei {k-m). Each of the four nuclei becomes the center of a spo-
roblast which secretes a membrane and transforms itself into a
spore (r?). Its nucleus, in the meantime, undergoes a division, and
two sporozoites develop in the spore (o). Oocysts leave the host in
crab gut, each liberating 3 sporozoites (6) which grow and produce
merozoites (lO/x by 2^) by schizogony in peri-intestinal connec-
tive tissue cells (6 chromosomes) (c-/) when host crab is eaten by
;
COCCIDIA 467
division (n, o); repeated nuclear division (p) forms many sporo-
blasts (q), each transforms itself into a spherical spore wdth 3 sporo-
zoites (Dobell; Naville; Belaf).
468 PROTOZOOLOGY
Genus Merocystis Dakin. Spoiogony in the kidney of the whelk,
Buccinum; schizogon}- unknown, in another host (possibh- a crab);
microgametocj^tes produce first cytomeres which in turn form
microgametes anisogamy gives rise to zygotes, zygote forms many
;
E. prevoti (Laveran and Mesnil) (Fig. 219, /). In the gut epithelium
of frogs; oocysts about IT/x by 12/x; when sporozoites are fully
formed, the spore membranes dissolve.
E. ranae Dobell (Fig. 219, j). In gut of frogs; oocysts 22jix bj'^ ISfx.
E. sardinae (Thelohan) (E. oxyspora Dobell) (Fig. 219, k). In
the testis of sardine; oocysts spherical 30-50^.
E. dupearum (Thelohan) {E. wenyoni Dobell) (Fig. 219, I). In the
hver of herrings, mackerels, and sprats; oocysts, spherical, 18-
33)U in diameter.
E. gadi Fiebiger. In the s\vhn-l)ladder cf (kulus I'irens, G. morrhua,
and 0. and sporogony; germination of spores
aeglefinus; schizogony
takes place in the bladder of the same host individual, bringing
about a very heavy infection; ooc3^sts 26-28^; pathogenic (Fiebiger).
Genus Jarfina Leger and Hesse. Oocysts ovoid, one end rounded
and the other drawn out into a shoit neck; 4 spores, each with 2
sporozoites.
/. paludom L. and H. (P ig. 219, m, n). In the gut cf Fulica atra and
Gallinula chloropus; ooc3'sts 15/i by llfx; sporulation in 15 days.
Genus Wenyonella Hoare. Oocj'sts with 4 spores, each with 4
sporozoites. Three species.
W. africana H. (Fig. 219, o, p). In the small intestine cf Boaedon
lineatus ("brown snake") Uganda; oocysts ovoid or subspherical,
in
18.5-19.2/x by 16-17.6^; spores ovoid, 9.6/i by 8m; sporulation in
5-6 days.
Genus Isospora Schneider. Oocyst produces 2 spores, each con-
taining 4 sporozoites.
7. hominis (Rivolta) (Figs. 219, q, r; 220). This is the sole coccidian
parasite of man known up to the present time. Its life cycle is un-
known, but most probably the schizogony, gametogenesis and sexual
fusion occur in the intestinal epithelium. Oocysts have only been
seen in the stools of infected persons.
The oocyst is asymmetrically' fusiform; 20-33^ by 10-16^; wall is
made up of two membranes which are highly resistant to chemicals;
when voided in faeces, the contents either fill up the oocyst or appear
as a spherical mass, composed of refractile granules of various sizes;
474 PROTOZOOLOGY
nucleus appears as a clear circular area; when the faecal specimen
is kept in a covered container at the room temperature, the proto-
plasmic mass divides into 2 spherical sporo blasts in about 24 hours
(Fig. 220, 3) each sporoblast develops in another 24 hours into a
spore (10-16m by T-lO/z) containing 4 sporozoites (4). Further
changes take place when the oocyst finds its way into the human in-
contaminated food or water.
testine in
hominis has been observed in widely separated regions, but
/.
incubation period was about six daA-s, the onset sudden, and the
duration over a month. The cure was spontaneous. The symptoms
were diarrhoea, abdominal discomfort, flatulence, lassitude, and loss
of weight. During the first three weeks of the illness no oocysts were
found, but then oocysts appeared in the stools for nine days. On the
10th day they were not seen, but reappeared on the 11th and 12th
days, after which the}' were not found again. The acute signs of ill-
ness abated within one week of the finding of the oocysts. The faeces
contained a large amount of undigested material, particularly fat
which gave it a thick oily consistency, showing signs of slow gaseous
formation. 7. hominis is to be considered as pathogenic to man
(coccidiosis) but although it gives rise to illness in some cases, the
;
10-14m by 7-9fi.
COCCIDIA 475
/. rivolta (Grassi) (Fig. 219, t). In the gut of cat and dogs; oocysts
20-25m by 15-20^.
I. felis Wenyon (Fig. 221, a). In cat and dog; oocysts 39-48^ by
26-37m.
I. suis Biester. In swine faeces; oocysts siibspherical, about
22.5/i by 19.4^t; sporulation in 4 days.
I. lacazei Labbe. In small intestine of passerine birds (sparrows,
blackbirds, finches, etc.); oocysts subspherical, 18.5-30^ by 18-
29.2)u; heavily infected sparrow shows definite symptoms; sporula-
tion in 4-5 days.
/. lieberkuhni (Labbe) (Fig. 221, h). Oocyst about 40m long; in the
kidney of frogs.
Genus Cyclospora Schneider. Development similar to that of
Eimeria; oocyst with 2 spores, each with 2 sporozoites and covM'ed
by a bi-valve shell.
Schaudinn (Fig. 221, c). In the gut of mole; sporo-
C. caryolyfica
zoites enter and develop in the nuclei of gut epithelial cells; oocyst
oval, about 15m by 11.5^l.
Genus Dorisiella Ray. Zygote develops (without becoming oocyst)
into 2 spores, each with 8 sporozoites; macrogametocytes migratory.
D. scolelepidis R. (Fig. 221, ci). In the gut of Scolelepis fuUginosa;
zygote contents divide into 2 oval spores, 12-16m by 6-10^; spore
with 8 sporozoites.
Genus Caryospora Leger. Oocyst develops into a single spore
with 8 sporozoites and a residual mass; membrane thick and yellow.
One species.
C. simplex L. (Fig. 221, e,f). In the gut-epithelium of Vipera aspis;
< cyst thick-walled, 10-15^ in diameter.
Genus Cryptosporidium Tyzzer. Lumen-dwelling minute organ-
ms; oocyst with 4 sporozoites.
C. muris T. (Fig. 221, g, i). In the peptic glands of the mouse; both
schizogony and sporogony in the mucoid material on surface of the
epithelium; oocysts 7m by 5m; 4 sporozoites, 12-14m long.
C. parvum T. In the glands of small intestine of the mouse; oocysts
with 4 sporozoites, 4.5m in diameter.
Genus Pfeififerinella Wasielewski. Macrogamete with a "recep-
tion tubule" by which microgamete enters; oocyst produces directly
8 sporozoites.
P. ellipsoides W. (Fig. 221, j). In the liver of Planorhis corneus;
oocysts oval, 13-15m long.
P. impudica Leger and Hollande (Fig. 221, k). In the liver of
Limax marginafiis; oocysts ovoid, 20m by 10m.
476 PROTOZOOLOGY
Genus Lankesterella Labbe. Oocyst produces 32 or more sporo-
zoites directly without spore-formation; in endothelial cells of cold-
Ijlooded vertebrates; mature sporozoites enter erythrocytes in which
they are transmitted to a new host individual by bloodsucking in-
vertebrates.
L. minima (Chaussat) (Fig. 221, /). In frogs; transmitted by
the leech (Placobdella marginata); frog acquires infection through
introduction of sporozoites by a leech; sporozoites make their way
into the blood capillaries of various organs; there they enter endo-
thelial cells;schizogony produces numerous merozoites which bring
about infection of many host cells; finally macro- and micro-gameto-
C3^tes are formed; anisogamy produces zygotes which transform into
oocysts, in which a number of sporozoites develop; these sporozoites
COCCIDIA 477
are set free upon disintegration of cyst wall in the blood plasma and
enter erythrocytes (Xoller) oocyst oval, about 33^1 by 23^.
;
Family 1 Adeleidae
In cells of circulatory system of vertebrates
Family 2 Haemogregarinidae (p. 480)
Fig. 222. The life-cycle of Adelea ovafa, X600 (Schellack and Reiche-
now). a, schizont entering the gut epithelium of the host centipede; b-d,
schizogony; e, larger form of merozoite; f, microgametocyte (left) and
macrogametocyte (right); g, association of gametocytes; h, i, fertilization;
j, zygote; k, nuclear division in zygote; 1, mature oocyst with many
spores.
480 PROTOZOOLOGY
Genus Klossiella Smith and Johnson. Microgametocyte produces
2 microgametes; oocyst with many spores, each with numerous
sporozoites; in the kidney of mammals.
K. muris S. and J. (Fig. 223, d, e). Oocyst with 12-16 spores;
spore with about 25 sporozoites, discharged in the host's urine; in the
kidney of mouse.
K. cobayae Seidelin. Oocyst with 8-20 spores; spore with about
30 sporozoites; in the kidney of guinea pig.
Genus Legerella Mesnil. Oocyst contains numerous sporozoites;
spores entirely lacking; in arthropods.
L. hydropori Vincent (Fig. 223,/). In the epithelium of Malpighian
tubules of Hydroporus palustris; oocysts ovoid, 20-25/x long, with
16 sporozoites which measure IT^t by 3/i.
References
THEthat
development
to
the Haemosporidia
the Coccidia
of
of
in
on the whole,
is,
484
HAEMOSPORIDIA 485
zoites upon entering the blood vessel, penetrate and enter immedi-
ately the erythrocyte and begin intracorpuscular development,
which process Schaudinn (1902-1903) reported to have seen in life.
Although some authors still follow this view, thei'e are increasing
numbers of others who doubt Schaudinn's observation, since no one
has up to the present time been able to duplicate the observation.
James (1931) noticed the ineffectiveness of quinine as a causal pro-
phylactic in malaria infection, and suggested the possibility that
the sporozoites are carried away immediately from peripheral to
visceral circulation and develop in the cells of reticulo-endothelial
system.
Boyd and Stratman-Thomas (1934) found that the peripheral
blood of a person who had been subjected to the bites of 15 anophe-
line mosquitoes infected by Plasmodium vivax, did not become in-
fectious to other per.sons by subinoculation until the 9th day and that
the parasites were not observed before the 11th day in the stained
films of the peripheral blood. Warren and Coggeshall (1937) observed
that when suspensions of the sporozoites of P. cathemerium ob-
tained from infected Culex pipiens, were inoculated into canaries,
the blood was not infectious for 72 hours, but emulsions made from
the spleen, liver and bonemarrow contained infectious parasites
which brought about infection by subinoculations in other birds.
These and many similar observations cannot be satisfactorily ex-
plained if one follows Schaudinn's view. The fact that P. elongatum
is capable of undergoing schizogony in the leucocytes and reticulo-
HAEMOSPORIDIA 487
g h
Fig. 226. Exoerythrocytic schizogony in avian Plasmodium, a-f, P.
gallinaceum in smears from chicks (James and Tate), a, monocyte from
lung, infected by 2 young schizonts; b, monocyte from liver, with a grow-
ing trinucleate schizont; c, monocyte from lung, with a large multi-
nucleate schizont; d, large mature schizont containing many mature mero-
zoites, free in lung; e, portion of broken schizont from lung, showing the
attached developing merozoites. ( X1660). f, a capillary of brain blocked
by 3 large schizonts ( X740). g, h, P. cathemerium in sections of organs of
mosquito
days; in P. malariae, 4-5 weeks, with the onset of fever lagging 3-12
days behind; and in two strains of P. falciparum, one, 6-25 days and
the other, 9-13 days; in another observation, P. falciparum was
observable in the peripheral blood in 5-9 days and the onset of fever
in 7-12 days.
The paroxysm of malaria is usually divisible into three stages: chill
P. vivax (Grassi and Feletti) (Fig. 228). The benign tertian malaria
parasite; schizogony completed in 48 hours and paroxysm every
third day. Ring forms: About 1/4-1/3 the diameter of erythrocytes;
unevenly narrow cytoplasmic ring is stained light blue (in Giemsa)
and encloses a vacuole; nucleus stained dark-red, conspicuous.
Growth period: Irregular amoeboid forms; host cell slightly' enlarged;
Schiiffner's dots begin to appear. Grown schizonts In about 26 hours
:
a
HAEMOSPORIDIA 493
cytes stain blue and contain a compact nucleus and coarser granules,
grouped around nucleus; microgametccj'tes stain less deeply blue or
reddish, and contain a large lightly staining nucleus and scattered
smaller haemozoin granules.
#
^ g h
f J
" ^m
W' ^^- i
e f g
# h
Fig. 231. Plasmodium ovale, X1535 (Original), a, ring-form; b, c,
growing schizonts; d-f, schizogonic stages; g, macroganietocyte; h. micro-
gametoeyte.
496 PROTOZOOLOGY
Table 9. Differential diagnosis of three species of human Plasmodium
HAEMOSPORIDIA 497
nucleus.
P. circumflexum Kikuth (Fig. 232, /). In red-winged blackbird,
cowbird and several other birds; growing schizonts and gametocytes
form broken rings around the host-cell nucleus; schizogony com-
pleted in 48 hours; 13-30 merozoites from a schizont; gametocytes
elongate, with a few haemozoin granules; transmitted by Culex and
Theobaldia.
P. polare Man well (Fig. 232, g). In cliff swallow {Petrochelidon I.
498 PROTOZOOLOGY
naevius) 8 merozoites from a schizont
; ; body outlines irregular, rough
gametocj'tes elongate.
l). In fire-back pheasant (Loph-
P. lophurae Coggeshall (Fig. 232,
ura i. from Borneo, examined at New York Zoological Park;
igniti)
8-18 merozoites from a schizont; gametoc^-tes large, elongate; host-
®®®(
cell nucleus not displaced; chicks are susceptible, but canaries re-
fractory. Experimentally it developed up to the oo cyst-stage in
Anopheles quadrimaculatus, though the sporozoites did not develop
(Coggeshall, 1941).
Mathis and Leger (1910) described this species from the teal duck
(Querquedula crecca) in Tonkin, China. Wick ware (1915) saw L.
anatis in ducks in Canada. O'Roke (1934) carried on experimental
studies on the developmental cycle with the form which he found in
wild and domestic ducks in Michigan. Herman (1938) observed the
502 PROTOZOOLOGY
According to O'Roke, the vector is a black fly, Simulium venustum,
in which the sexual reproduction takes place. Gametocytes develop
into mature gametes in 1-2 minutes after blood is obtained from an
infected duck; macrogametes about 8m in diameter; 4-8 micro-
gametes, 15.7-24. 1/i long, from a single microgametocyte; zygotes
are found in stomach contents of fly in 10-20 minutes after sucking
in the infected blood cf bird; motile ookinetes abundant after 5 hours,
measure 33.3)u by 3-4. 6m; 22 hours after sucking duck blood, oocysts
are found on outer wall of stcmach; sporozoites mature probably in
24-48 hours; 5 days after a duck has been bitten by infected black
flies, schizogonic stages are noticed in endothelial cells of capillaries
of lungs, liver, spleen;on about 7th day gametocytes appear in blood
liver and spleen become hypertrophied; the infection among duck-
lings is said to be highly fatal and appears often suddenly.
Mathis and Leger: Macrogametocytes, oval; 14-15m by 4.5-5.5m;
several vacuoles in darkly stained cytoplasm. Microgametocytes,
oval; slightly smaller; cytoplasm stains less deeply. Infected host
cellsabout 48m long; nucleus elongate.
Huff found that (1) young schizonts are in macrophages of, and
also extracellularly in, the spleen and hver; (2) two types of schi-
zonts occur: one, "hepatic schizonts" in hepatic cells which cause
no distortion or alteration of the host cell, and the other, "megalo-
schizonts" in the blood vessels of, or extravascularly in, the heart,
spleen, hver and intestine; (3) megaloschizonts become divided into
many cytomeres which give rise to numerous merozoites; (4) young
gametocytes occur in lymphocytes, monocytes, myelocytes and late
polychromatophile erythrc blasts; (5) the cells in which fully grown
gametocytes occur, appear to be macrophages.
^ %^ ^ ^
Fig. 237. Toxoplasma gondii. X about 1750. (Chatton and Blanc)
a, isolated organisms; b, 2 trophozoites; c,organisms undergoing binary
fission; d, a host cell with many organisms which developed by repeated
binary fission.
References
Boyd, M. F. 1930 An
introduction to malariology. Cambridge.
1940 On strains or races of the malaria parasites. Amer.
Jour. Trop. Med., Vol. 20.
and W. K. Stratman-Thomas 1933 A controlled technique
for the employment of naturally induced malaria in the therapy
of paresis. Amer. Jour. Hyg., Vol. 17.
Chen, T. T. 1944 The nuclei in avian malaria parasites. I. The
structure of nuclei in Plasmodium elongatum, with some con-
siderations on technique. Amer. Jour. Hyg., Vol. 40.
CoATNEY, G. R. and R. L. Roudabush 1937 Some blood parasites
from Nebraska birds. Amer. Midi. Nat., Vol. 18.
Dennis, E. W. 1932 The life-cycle of Babesia higemina (Smith and
Kilbourne) of Texas cattle-fever in the tick, Margaropus an-
nulatus (Say). Uni. Calif. Publ. Zool., Vol. 36.
Hartman, E. 1927 Three species of bird malaria. Arch. f. Pro-
tistenk., Vol. 60.
Hewitt, R. 1940 Bird malaria. Amer. Jour. Hyg., Monogr. Ser.,
No. 15.
Huff, C. G. 1942 Schizogony and gametocyte development in
Leucocytozoon simondi, and comparison with Plasmodium and
Haemoproteus. Jour. Infect. Dis., Vol. 71.
James, S. P. and P. Tate 1938 Exo-erythrocytic schizogony in
Plasmodium gallinaceum Brumpt, 1935. Parasitology, Vol. 30.
O'RoKE, E. C. 1934 A malaria-like disease of ducks caused by
Leucocytozoon anatis Wickware. Uni. Michigan Sch. Forest and
Conservation, Bui., No. 4.
Porter, R. J. and C. G. Huff 1940 Review of the literature on
exo-erythrocytic schizogony in certain malarial parasites and
its relation to the schizogonic cycle in Plasmodium elongatum.
Amer. Jour. Trop. Med., Vol. 20.
Ross, R. 1928 Studies on malaria. London.
Sabin, A. B. Toxoplasmosis. A recently recognized disease of hu-
man beings. In: Advances in pediatrics, edited by A. G. De Sanc-
tis. Vol. 1.
Simmons, J. S. et al. 1939 Malaria in Panama. Amer. Jour. Hyg.,
Monogr. Ser. No. 13.
Stratman-Thomas, W. K. 1940 The influence of temperature on
Plasmodium vivax. Amer. Jour. Trop. Med., Vol. 20.
Wenyon, C. M. 1926 Protozoology. Vol. 2. London and Baltimore.
Chapter 27
Subclass 2 Acnidosporidia Cepede
508 PROTOZOOLOGY
may vary in size from microscopic to as large as 5 centimeters. They
are cylindrical with more or less pointed extremities and with a some-
what lobulated surface, and appear opaque whitish. They were for-
merly called Miescher's tubes (Fig. 238). The envelope around the
parasitic mass appears to rupture sooner or later and the spores are
set free in the blood stream and into the alimentary canal. The spores
find finally their way out of the host intestine and become the source
of infection (Scott).
Muscle layer
Fibrous zone
Externals
Median / Cyst membrane
Internal
Sporoblasts
Spores
<^%^
%
^
^^» ^<v;>.
St. Remy) ;
parasites 84^ by 27)Lt and spores 4.25m by 1.75m (Darling)
parasites spherical, 500m in diameter and spores over 10m long
(Manifold); parasites 5.3 cm. by 320m and spores 8.33m by 1.6m
(Vasudevan). The parasitic masses are oval to spindle in form and
imbedded in the muscle cells which are distended, and may appear
white-streaked to naked eye. Seen in sections, the body is divided
into compartments. Gilmore, Kean and Posey (1942) have recently
found three bodies in sectioned heart muscles of an eleven year old
child who died from an unknown cause, and considered them as
sarcosporidian bodies. They measured 25m by 19m, 57m by 30m, and
510 PROTOZOOLOGY
41m by 25m in cross-sections; there were no septa within the bodies;
minute bodies present in the masses were mostly rounded and about
l/i in diameter, though a few were crescentic. The questions such as
what species infect man, how man becomes infected, etc., are un-
answered at present.
*S. tenella Railliet (Figs. 238, a; 239). In the muscles of tongue,
pharynx, oesophagus, larynx, neck, heart, etc., of sheep; large
parasites 40/x-2 cm. long with a thin membrane; spores sickleform.
S. miescheriana (Kiihn) (Fig. 238, b). In muscles of pig; parasitic
mass up to 3-4 mm. by 3 mm; envelope striated; spores reniform,
capable of division when young (Manz).
S. bertrami Doflein. In the muscles of horse; similar to S. miescher-
iana; parasitic mass up to 9-10 mm. envelope striated.
;
become paired (J, g) and the nuclear membranes disappear ih). The
which develops into a spore; spore truncate with a lid at one end;
envelope sometimes prolonged into processes; in aquatic annelids
and molluscs.
H. chitonis (Lankester) (Fig. 242, a, b). In liver and connective
tissue of Craspidochilus cinereus; spores oval, 10/* by 6m; envelope
with 2 prolonged projections.
H. limnodrili Granata (Fig. 242, c). In gut epithelium of Lim-
nodrilus udekemianus; spores 10-12^ by 8-10/x.
512 PROTOZOOLOGY
H. nemertis Debaisieux (Fig. 242, d). In connective tissue of
Lineus hilineatus; spores oval with a fiat operculum, but without
any projections of envelope, 7m by 4^.
H. heterocirri C. and M. (Fig. 242, e). In gut epithelium of Het-
erocirrus viridis; mature organisms 50-60m by 30-40m; spores 6.5m
by 4m.
H. scolopli C. and M. (Fig. 242, /). In Scoloplos millleri; fully
grown form 100-150m by 20-30m; spores 10m by 6.5m.
THE members
ant spores which
of this subclass possess without exception resist-
are of unique structure. Each spore possesses
1-4 polar filaments and one to many sporoplasms. The membrane
which envelops these structures may be a single-piece or bi- or tri-
valved. The polar filament is typically coiled within a polar capsule.
In the order Myxosporidia and Actinomyxidia, there appear
several cells during the process of sporulation. These cells give rise
to one to many sporoplasms or generative cells, capsulogenous
cells, and spore membrane. This condition is not observed in other
groups of Protozoa and for this reason some writers recognize a close
between these two orders and the Mesozoa. The method of
affinity
multiplication in the Cnidosporidia is schizogonic and sporogonic.
515
516 PROTOZOOLOGY
filament, varies in number from one to four. Except in the family
Myxidiidae, in which one polar capsule is situated near each of the
poles of the spore, the polar capusles are always grouped at one end
which is ordinarily designated as the anterior end of the spore. Below
or between (in Myxidiidae) the polar capsules, there is almost always
a sporoplasm. Ordinarily a young spore possesses two sporoplasm
nuclei which fuse into one (autogamy) when the spore becomes
mature. In Myxobolidae there is a glycogenous substance in a vacu-
ole which stains mahogany red with iodine and is known as the
iodinophilous (iodophile) vacuole.
The Myxosporidia are exclusively of lower verte-
parasites
brates, especially fishes. Both fresh water fishes have been
and salt
found to harbor, or to be infected by, Myxosporidia in various
regions of the world. A few occur in Amphibia and Reptilia, but no
species has been found to occur in either birds or mammals. When
a spore gains entrance into the digestive tract of a specific host fish,
the sporoplasm leaves the spore as an amoebula which penetrates
through the gut-epithelium and, after a period of migration, enters
CNIDOSPORIDIA, MYXOSPORIDIA 517
expense of the host tissue cells, and the nucleus divides repeatedly.
Some nuclei become surrounded by masses of dense cytoplasm
and become the sporonts (Fig. 243). The sporonts grow and their
nuclei divide several times, forming 6-18 daughter nuclei, each with
a small mass of cytoplasm. The number of the nuclei thus produced
depends upon the structure of the mature spore, and also upon
whether 1 or 2 spores develop in a sporont. When the sporont de-
velops into a single spore, it is called a monosporoblastic sporont,
and if two spores are formed within a sporont, which is usually the
case, the sporont is called disporoblastic, or pansporoblast. The
spore-formation begins usually in the central area of the large tro-
phozoite, which continues to grow. The surrounding host tissue
becomes degenerated or modified and forms an envelope that is
often large enough to be visible to the naked eye. This is ordinarily
referred to as a myxosporidian cyst. If the site of infection is near the
body surface, the large cyst breaks and the mature spores become
set free in the water. In case the infection is confined to internal
organs, the spores will not be set free while the host fish lives. Upon
its death and disintegration of the body, however, the liberated
become the source of new infection.
spores
The more primitive Myxosporidia are coelozoic in the host's
organs, such as the gall bladder, uriniferous tubules of the kidney,
urinary bladder, etc. In these forms, the liberated amoebulae make
their way into the specific organ and there grow into multinucleate
amoeboid trophozoites which are capable of forming pseudopodia
They multiply by exogenous or endogenous bud-
of various types.
ding or plasmotomy. One to several spores are developed in the
trophozoite.
Almost all observers agree in maintaining the view that the 2
nuclei of the sporoplasm or 2 uninucleate sporoplasms fuse into one
(autogamy or paedogamy), but as to the nuclear as well as cyto-
plasmic changes priorto, and during, spore-formation, there is a
diversity of opinions. To illustrate the views held by those who be-
lieve there is a sexual phase in the development of a myxosporidian,
Sphaeromyxa sabrazesi may be taken as an example. De-
(p. 528)
baisieux's observation on
myxosporidian is in brief as follows
this
(Fig. 244) Sporoplasms after finding their way into the gall bladder
:
of host fish develop into large trophozoites containing many nuclei (a),
2 vegetative nuclei become surrounded by a cytoplasmic mass (c)
and this develops into a primary propagative cell (d) which divides
(3 chromosomes are noted) (e) and forms secondary propagative
518 PROTOZOOLOGY
cells (/). A binucleate sporocyte is formed from the latter by unequal
nuclear division ig-i) and 2 sporocytes unite to form a tetranucleate
pansporoblast ij) which develops into 2 spores (A;, I). Sporoplasm
shows first 2 nuclei, but later 4, of which 2 degenerate and the other 2
fuse into one nucleus. On the other hand, according to Naville (1930)
Xi (Kudo).
fishes, while the gall bladder and urinary bladder of marine fishes
harbor one or more species of Myxosporidia. When the infection is
concentrated in the fins or integument, the resulting changes are
quite conspicuous (Fig. 246). The infection in the gills is usually
manifest by whitish pustules which can be frequently detected with
the unaided eye. When the wall of the alimentary canal, mesentery,
liver, and other organs are attacked, one sees considerable changes
in them. Heavy myxosporidian infection of the gall bladder or uri-
nary bladder of the host fish may cause abnormal appearance and
coloration or unusual enlargement of the organ, but under ordinary
circumstances the infection is detected only by a microscopical ex-
Eurysporea
Suborder 1
Family 1 Ceratomyxidae
Spores less laterally expanded; in freshwater fish; histozoic or coelozoic. .
:< .';,.:1?**M;'. • ;
-f i
Family 2 Triactinomyxidae
Genus Triactinomyxon Stole. Each of 3 shell-valves drawn out
into a long process, the whole anchor-like; spore with 8 or more
uninucleate sporoplasms; in the gut-epithelium of oligochaetes.
T. ignotum S. (Fig. 254, d). Spore with 8 sporoplasms; in Tuhifex
tubifex.
532 PROTOZOOLOGY
T. magnum Granata. Spore with 16 sporoplasms; in Limnodrilus
udekemianus.
T. legeri Mackinnon and Adams. Spore with 24 sporoplasms; in
Tubifex tubifex.
T. diihium G. Spore with 32 sporoplasms; in Tubifex tubifex.
References
AuERBACH, M. 1910 Die Cnidosporidien. Leipzig.
Caullery, M. and F. Mesnil 1905 Recherches sur les Actinomy-
Arch. f. Protistenk., Vol. 6.
xidies.
Davis, H. S. 1917 The Myxosporidia of the Beaufort region. Bull.
U. S. Bureau Fish., Vol. 35.
Fantham, H. B., a. Porter and L. R. Richardson 1939 Some
Myxosporidia found in certain fresh-water fishes in Quebec
Province, Canada. Parasitology, Vol. 31.
1940 Some more Myxosporidia observed in
Canadian fi,shes. Ibid., Vol. 32.
Granata, L. 1924 Gli Attinomissidi. Arch. f. Protistenk., Vol. 50.
534 PROTOZOOLOGY
Kudo, R. R. 1920 Studies on Myxosporidia. Illinois Biol. Monogr.,
Vol. 5.
1933 A taxonomic consideration of Myxosporidia. Trans.
Amer. Micr. Soc, Vol. 52.
1934 Studies on some protozoan parasites of fishes of Illi-
Monogr., Vol. 32.
nois. Illinois Biol.
1943 Further observations on the protozoan, Myxidium
serotinum, inhabiting the gall bladder of North American Salien-
tia. Jour. Morph., Vol. 72.
1944 The morphology and development of Nosema nota-
hilis Kudo, and of its host, Sphaerospora polymorphja Davis,
parasitic in Opsanus tau and 0. beta. Illinois Biol. Monogr.,
Vol. 20.
Naville, a. 1930 Le cycle chromosomique d'une nouvelle Actino-
mvxidie: Guyenotia sphaerulosa n.gen., n.sp. Quart. Jour. Micr.
Sci., Vol. 73.
Chapter 29
Order 3 Microsporidia Balbiani
protozoan parasites.
trophied nucleus of adipose tissue of larval Ci lex pipiens, the latter due to
infection by Stempellia magna, XlOOO (Kudo).
535
536 PROTOZOOLOGY
host (Fig, 256), the polar filaments are extruded and perhaps anchor
the spores to the gut-epithelium (a). The sporoplasms emerge
through the opening after the filaments become completely de-
tached (b). By amoeboid movements they penetrate through the in-
testinal epithelium and enter the blood stream or body cavity and
reach the specific site of infection (c). They then enter the host cells
and undergo multiplication at the expense of the latter (d-n).The
trophozoites become sporonts, each of which produces a number of
6" 0^ B^
0-6 oo
^
K m
(
Fig. 260. a, b, Pyrotheca incurvata, X2000 (Hesse); c, d, Coccospora
slavinae (d, with extruded polar filament), X2000 (L^ger and Hesse);
e, f, Mrazekia caudata (e, an infected host cell, X700 (Mrazek) f, a spore,;
cilia
which serve as cell-organs of loco-
are present only during early devel-
opmental stages. The members of this subphylum possess a unique
organization not seen in the Plasmodroma; namely, except Proto-
the Ciliophora contain two kinds of nuclei: the macronucleus
ciliata,
and the micronucleus. The former is large and massive, and controls
the metabolic activities of the organism, while the latter is minute
and usually vesicular or less compact, and is concerned with the
reproductive processes. Nutrition is holozoic or parasitic; holophytic
in Cyclotrichium meunieri (p. 565). Sexual reproduction is mainly
by conjugation, and asexual reproduction is by binary fission or
budding. The majority are free-living, but a number of parasitic
forms also occur.
The Ciliophora are divided into two classes:
545
546 PROTOZOOLOGY
food vacuoles, contractile vacuoles, pigment granules, crystals, etc.
In the majority of ciliates, the anterior and posterior extremities are
permanent and distinct; in all cytostome-possessing forms, the oral
and aboral surfaces are distinguishable, while in numerous creeping
forms the dorsal and ventral sides are differentiated.
The body is covered by a very thin yet definite membrane, the
pellicle, which is ordinarily uniformly thin and covers the entire
be closed when the animal is not feeding. The cytostome opens into
the cytopharynx (or gullet), a canal which ends in the deeper portion
of the endoplasm. In the cytopharynx there may be present one or
more undulating membranes to facilitate intaking of the food. Oc-
casionally the cytostome is surrounded by trichites or trichocysts
(p. 62). When the cytostome is not at the anterior region as, for
instance, in Paramecium, there is a peristome (or oral groove) which
starts at or near the anterior end and runs posteriorly. The peristome
is ciliated so that food particles are thrown down along it and ulti-
mately into the cytostome which is located at its posterior end. Solid
waste particles are extruded from the cytopyge, or cell-anus, which
is usually noticeable only at the time of actual defecation (p. 92).
Subclass 1 Protociliata
Macronucleus and micronucleus; sexual reproduction conjugation
Subclass 2 Euciliata (p. 551)
References
BtJTscHLi, O. 1887-1889 Protozoa. In: Bronn's Klassen und Ord-
nungen des Thier-reichs. Vol. 1.
DoFLEiN, F. and E. Reichenow 1929 Lehrbuch der Protozoen-
kunde. Jena.
Kahl, a. 1930-1935 Urtiere oder Protozoa I: Wimpertiere oder
Ciliata (Infusoria). In Dahl: Die Tierwelt Deidschlands und der
angrenzenden Meeresteile nach ihren Merkmalen und nach ihrer
Lebensweise. Parts 18, 21, 25, 30.
Kent, W. S. 1880 to 1882 A manual of Infusoria.
Stein, F. 1867 Der Organismus der Infusionsthiere. Vol. 2.
Stokes, A. C
1888 A preliminary contribution toward a history of
the fresh-water Infusoria of the United States. Jour. Trenton.
Natural Hist. Soc, Vol. 1.
The members of this order show uniform ciliation over the entire
body surface. Adoral zone does not occur. The majority possess a
cytostome which varies among different forms. Nutrition is holo-
zoic or saprozoic. Asexual reproduction is usually by transverse
fission and sexual reproduction by conjugation. Encystment is com-
mon. The holotrichous ciliates are conspicuous free-living forms in all
sorts of fresh, brackish, and salt waters, though some are parasitic.
The order is here divided into 6 suborders:
551
552 PROTOZOOLOGY
Suborder 1 Astomata Schewiakoff
The ciliates placed in this suborder possess no cytostome, although
there may occur a slit-like organella which has been looked upon as
a vestigial cytostome. The body ciliation is usually uniform. Asexual
division is carried on by transverse fission and often by budding
which results in chain formation. Sexual reproduction is conjugation
and in some encystment is known. These organisms are parasitic in
various invertebrates living in fresh or salt water.
Several species.
H. secans S. Elongated; 160-500^ by 20-35^; 15-30 contractile
vacuoles in a row; spicule 10-15m long; in the intestine of Lwnhricus
variegatus.
Genus Radiophrya Rossolimo. Elongate, often with satellites;
558 PROTOZOOLOGY
R. hoplites R. 100-IOOOm long; in the intestine of
(Fig. 266, d, e).
Lamprodrilus, Teleuscolex, Styloscolex, and other oHgochaetes.
Genus Metaradiophrya Heidenreich. Ovoid to elhpsoid; with 2
rows of contractile vacuoles; with a hook attached to a long
lateral
shaft; ectoplasmic fibers supporting the hook; in the intestine of
oligochaetes. Several species.
References
Beers, C. D. 1938 Structure and division in the astomatous ciliate
Metaradiophrya asymmetrica n. sp. Jour. E. Mitchell Sci. Soc,
Vol. 54.
Cepede, C. 1910 Recherches sur les Infusoires astomes. Arch. zool.
exp., Vol. 3 (ser. 5).
Cheissin, E. 1930 Morphologic und systematische Studien iiber
Astomata aus dem Baikalsee. Arch. f. Protistenk., Vol. 70.
Heidenreich, E. 1935 Untersuchungen an parasitischen Ciliaten
aus Anneliden. I, II. Arch. f. Protistenk., Vol. 84.
MacLennan, R. F. 1944 The pulsatory cycle of the contractile
canal in the ciliate Haptophrya. Trans. Amer. Micr. Soc, Vol.
63.
Rossolimo, L. L. 1926 Parasitische Infusorien aus dem Baikalsee.
Arch. f. Protistenk., Vol. 54.
Chapter 32
Order 1 Holotricha Stein (continued)
560
HOLOTRICHA 561
end, with 4 rows of long cilia; body surface with shorter cilia;
its lip
Vasicola; sapropelic.
P. (Vasicola) grandis (Penard) (Fig. 268, d). Free-swimming;
elongated fusiform; numerous contractile vacuoles on one side; body
125-220^1 long; sapropelic in fresh water.
M. acarus Stein (Fig. 268, I, m). Oral tentacles with capitate tip;
10-16m long; salt water, Florida (Noland).
Genus Askenasia Blochmann. Resembles Didinium; ovoid; with
HOLOTRICHA 565
formed after dropping off the fish skin and in which numerous (up to
1000) ciliated bodies (30-45^ in diameter) are produced; conjugation
has been reported; parasitic in the integument of freshwater fishes;
in aquarium, host fish may suffer death; widely distributed.
tion coarse and not over entire body surface; resembles somewhat
Coleps; fresh water.
P. ovatus K. (Fig. 272, h). Caudal bristle breaks off easily; body
30m long; fresh water among decaying vegetation.
Genus Rhopalophrya Kahl. Cylindrical; furrows widely separated;
slightly asymmetrical; curved ventrally; dorsal surface convex;
ventral surface flat or slightly concave; anterior end with 'neck'; 2
spherical macro nuclei fresh or
; salt water; sapropelic.
R. salina Kirby (Fig. 273, a). Cylindrical, tapering gradually to a
truncated anterior end, slightly curved ventrally; cilia (6-10^ long)
sparsely distributed; 2 macro nuclei, spherical; 29-55ju long; 16-21)u
in diameter; in concentrated brine (salts "34.8 per cent"; pH 9.48)
from Searles Lake; California.
Genus Enchelyodon Claparede and Lachmann. Elongated; cy-
lindrical, ovoid or flask-shaped; some with head-like prolongation;
cyto pharynx with trichites; cilia long at anterior end; fresh or salt
contracted pellicular striae not spiral and no neck as is the case with
Chaenea; salt water. Many species.
T. phoenicopterus Cohn (Fig. 273, d, e). Elongate; extensible and
contractile; neck and tail distinct when contracted; cytostome at
anterior end, surrounded by a ridge containing indistinctly visible
short trichocysts, cytopharynx with trichocysts; macronuclei made
up of 4 radially arranged endosomes suspended in the nucleoplasm
(Gruber, KahJ); micronucleus difficult to make out; contractile
terus (d, XlOO; e, anterior end, X220) (Kahl); f, g, T. subviridis (f, Xl30;
g, nucleus, X400) (Noland); h, Parachaenia myae, X670 (Kofoid and
Bush).
References
HsiUNG, T. S. 1930 A monograph on the Protozoa of the large in-
testine of the horse. Iowa St. Coll. Jour. Sci., Vol. 4.
Kahl, a. 1930 In Dahl's Die Tierwelt Deutschlands. Part 18.
KiRBY Jr., H. 1934 Some ciUates from salt marshes in California.
Arch. Protistenk., Vol. 82.
f.
line; with flattened neck and tail, both of which are moderately
contractile; posterior end bluntly rounded; without trichocysts;
neck stout, bent toward the dorsal side; cytostome a long slit; con-
tractile vacuole posterior; 2 spherical macronuclei between which a
micronucleus is located; 100m long; fresh water and probably also in
salt water.
Genus Loxophyllum Dujardin {Opisthodon Stein). Generally simi-
lar to Lionotus in appearance; but ventral side with a hyaline border,
580
HOLOTRICHA 581
more micronuclei; 5-25 Miiller's vesicles (p. 76; Fig. 30, a, h) in dor-
sal region; fresh water.
L. vorax Stokes (Fig. 277, g). 125-140^ long; yellowish brown, a
row of slightly longer cilia; sapropelic in standing fresh water.
L. magnus S. (Fig. 277, h). Extended about 700^ long; dark brown;
12-20 or more Miiller's vesicles in a row along dorsal border; stand-
ing pond water.
Genus Remanella Kahl. Similar to Loxodes in general appearance;
but with endoskeleton consisting of 12-20^ long spindle-form needles
lying below broad ciliated surface in 3-5 longitudinal strings con-
nected with fibrils; Miiller's vesicles (Fig. 30, c) in some, said to be
different from those of Loxodes (Kahl); sandy shore of sea.
R. rugosa K. (Fig. 277, i, j). 200-300^ long.
HOLOTRICHA 585
.^\XU^^
ginning on each side near anterior end, united at the notched pos-
terior end; a series of apertures in grooves considered as cytostomes;
at posterior 1/3, an aperture gives rise to branching canals running
through endoplasm, and is considered as excretory in function; in
the colon of Procavia capensis and P. brucei. One species.
References
Chatton, E. and C. Perard 1921 Les Nicollelidae, infusoires in-
testinaux des gondis et des damans, et le 'cycle 6volutif des
Bull. biol. Fr. et Bel., Vol. 55.
cilies.'
Kahl, a. 1931 Urtiere oder Protozoa. Dahl's Die Tierwelt Deutsch-
lands. Part 21.
Kent, W. S. 1880-1882 A manual of Infusoria.
Stein, F. 1867 Der Organismus der Infusionstiere. Vol. 2.
Stokes, A. C. 1888 A preliminary contribution toward a history of
the freshwater Infusoria of the United States. Jour. Trenton
Nat. Hist. Soc, Vol. 1.
Wenrich, D. H. 1929 Observations on some freshwater ciliates.
II. Paradileptus, n.g. Trans. Amer. Micr. Soc, Vol. 48.
.
Chapter 34
Order 1 Holotricha Stein (continued)
tened; aboral surface convex; oral surface flat or concave; with a few
deep longitudinal furrows; ciliation sparse; cytostome and a small
cytopharynx simple, near the middle of body; fresh water. Several
species.
D. dentata F. (Fig. 281, g). With a small process near cytostome;
2 rows of ciliary furrows on both oral and aboral surfaces; cilia on
with densely standing short cilia; macro nucleus spherical; several (?)
micro nuclei; contractile vacuole flattened, terminal; in salt water.
W. rostrata K. (Fig. 285, i). 120-180ai long; salt water culture with
Oscillatoria.
W. nictabolica Johnson and Larson. 85-400iu long; division cysts
85-1 55/i in diameter; resting cysts 40-62/x in diameter; in freshwater
ponds. Johnson and Evans (1939, 1940) find two types of protective
cysts in this ciliate: "stable"and "unstable" cysts, formation of both
of which depends upon the absence of food. These cysts have three
membranes: a thin innermost endocyst, a rigid mesocyst and a gela-
tinous outer ectocyst. The protoplasmic mass of the stable cyst is
smaller, and free from vacuoles, and its ectocyst is thick, while that
of the unstable cyst is larger, contains at least one fluid vacuole and
its ectocyst is very thin. Crowding, feeding on starved Paramecium,
rod which is imbedded in the margin along the right wall of the oral
cavity and which he named stomato style.
Genus Biggaria Kahl. Scoop-like form; anterior 2/3 thin, posterior
region thickened, terminating in a rudder-like style; cilia in longi-
References
Johnson, W. H. and F. R. Evans 1939 A
of encystment in
study
the ciUate, Woodruffia metabolica. Arch.
Protistenk., Vol. 92.
f.
608
HOLOTRICHA 609
end and bluntly pointed narrow posterior end; preoral canal wide;
a dorsal ridge in posterior region of body; macro nucleus sausage-
form; a micro nucleus; contractile vacuole in middle of body, with
long collecting canals; in fresh water.
water.
T. vorax (Kidder, Lilly and Claff) {Glaucoma vorax K. L. and C.)
(Fig. 38). Form and size vary; bacteria-feeders elongate pyriform,
50-75)u long; sap ro zoic forms fusiform, 30-70/x long, decreasing in
size with the age of culture; sterile particle-feeders, 60-80^ long;
carnivores and cannibals broadly pyriform, 100-250^ long; 19-21
ciliary meridians; macronucleus ovoid, central; in carnivores, out-
line irregular; a single micronucleus; pond water.
Genus Leucophrys Ehrenberg. Broadly pyriform; cytostome large,
pyriform, with its axis parallel to body axis; ectoplasmic flange along
Fig. 289. Diagrammatic oral views and oral apparatus of four genera.
Ciliary meridians of: a, Tetrahymena geleii; b, Leucophrys patula; c, Glau-
coma scintillans; and d, Colpidium campijlum. All X about 535. e, cyto-
stome of Tetrahymena geleii, showing the ectoplasmic ridge and 3 mem-
branellae on left and a large undulating membrane on right. Diagrams
of cytostome of: f, T. geleii; g, Leucophrys patula; h, Glaucoma scintillans;
and i, Colpidium colpoda. (a, c, d, modified after Kidder; b, e-i, modified
after Furgason.)
tenth the body length 55-60 ciliary meridians; preoral suture curved
;
42-50/x long.
Genus Platynematum Kahl. Ovoid or ellipsoid; highly flattened;
with a long caudal cilium; contractile vacuole posterior-right; small
cytostome more or less toward right side, with 2 outer membranes;
ciliary furrows horseshoe-shaped; in fresh or salt water.
P. sociale (Penard) (Fig. 290, h). Anterior half more flattened;
ventral side concave; cytostome in the anterior third; yellowish and
granulated; 30-50jLt long; sapropelic in fresh and brackish water.
HOLOTRICHA 615
*S. muscorum Kahl (Fig. 290, j). Cytostome large, with a large
(Rossolimo).
tral side close to left border in the anterior half, with a small tongue-
like membrane; cyto pharynx indistinct; macro nucleus spherical,
central; contractile vacuole terminal; in salt or fresh water.
HOLOTRICHA 617
P. anodontae Kahl (Fig. 292, b). About 55^ long; posterior cilium
about 1/2 the body length; in Sphaerium, Anodonta.
P. setigerum Calkins (Fig. 292, c, d). Ellipsoid; flattened; ventral
surface slightly concave; about 25 ciliary rows; 38-50m long (No-
land); in salt water; Massachusetts, Florida.
P. coronatum Kent (Fig. 292, e). Elongate ovoid; both ends equally
rounded; caudal cilia long; about 40 ciliary rows; 47-75/1 long
(Noland); in fresh and salt water; Florida.
HOLOTRICHA 619
f
^^^/ d
1/6 the body length; a few cilia at posterior end; oval macronucleus
central; contractile vacuole posterior; about 60m long; in fresh water.
C. cacci Powers (Fig. 293, g). About 32-92^ long; in the intestine of
Tripneustes esculentus and other echinoids; Tortugas.
References
BuLLiNGTON, W. E. 1939 Astudy of spiraling in the ciliate Fron-
tonia with a review of the genus and a description of two new
species. Arch. f. Protistenk., Vol. 92.
BuRBANK, W. D. 1942 Physiology of the ciliate Colpidium colpoda.
I. The effect of various bacteria as food on the division rate of
Without tentacles
Ciliation uniform; ciliary rows meridional, close; peristome does not
begin near the anterior end
Thigmotactic cilia on entire broad side; with large peristome
Family 1 Conchophthiridae
623
624 PROTOZOOLOGY
cavity, gills and on non-ciliated surface of palps of Elliptio com-
planatus; Woods Hole.
C. curtus Engelmann. Somewhat broader; 60-125)U by 50-90)u;
peristomal smaller; cytopharynx less conspicuous, but longer;
field
ciliation dense; endoplasmic granules are more closely packed and
do not extend as far out toward anterior end; macro nucleus central.
Kidder found this ciliate in the mantle cavity of Anodonta marginata,
A. implicata, A. catarecta, Lampsilis radiata, L. cariosa and Alasmi-
donta undulata which were obtained from the freshwater lakes of
Massachusetts and New York.
C. magna Kidder. Much larger; 123-204/i by 63-116m; closer cilia-
tion; anterior 1/3 filled with smaller granules; irregularly outlined
macro nucleus, 25-30m in diameter, central; 2 (or 1) micro nuclei;
aperture for contractile vacuole large; mantle cavity of Elliptio com-
planatus; Massachusetts.
C. caryoclada K. (Fig. 294, h). Oval; extremely flattened; leaf-
like; cytostome small, in posterior fourth; macronucleus conspic-
uously branched; 2 (or 1) micronuclei; 140-250^ by 90-160m; mantle
cavity of the edible clam, Siliqua patula; Oregon.
C. mytili de Morgan (Fig. 55). Reniform; 130-220^ by 76-161^;
peristomal groove on the right side; trichocysts conspicuous along
frontal margin; macronucleus oval; 2 micronuclei. Kidder (1933)
found the organism on the foot of the common mussel, Mytilus
edulis, in New York and studied its division and conjugation.
Genus Myxophyllum Raabe. Oval or spheroid pellicle elastic and
;
since its affinity to other forms is not yet clear. Beers who placed it
626 PROTOZOOLOGY
Genus Ptychostomum Stein {Lada Vejdovsky). Sucker circular or
ovoid; macronucleus ovoid or reniform, not elongate; in oligochaetes.
Several species.
P. hacteriophilum Miyashita (Fig. 294, e). Elongate oval; 70-130/i
by 30-45m; sucker oval and large, about SO/x in diameter; macronu-
cleus ellipsoid ; endoplasm with numerous rods (symbiotic bacteria?)
in Criodrilus sp. (Oligochaeta).
References
Beers, C. D. 1938 Hysterocineta eiseniae n.sp., an endoparasitic
ciliate from the earthworm Eisenia lonnhergi. Arch. f. Protis-
tenk.. Vol. 9L
Chatton, E. and A. Lwoff 1926 Diagnoses de cilies thigmotriches
nouveaux. Bull. soc. zool. Fr., Vol. 51.
Cheissin, E. 1931 Infusorien Ancistridae und Boveriidae aus dem
Baikalsee. Arch. f. Protistenk., Vol. 73.
Kahl, a. 1931, 1935 Dahl's Die Tierwalt Deutschlands. Parts 21,
30.
Kidder, G. W. 1933 Studies on Conchophthirus mytili de Morgan.
Arch. f. Protistenk., Vol. 79.
I, II.
1933 On the genus Ancistruma Strand (Ancistrum Mau-
pas). I. Biol. Bull. Vol. 64; 11. Arch. f. Protistenk., Vol. 81.
1934 Studies on the ciliates from freshwater mussels. I, II.
Biol. Bull, Vol. 66.
MacLennan, R. F. and F. H. Connell 1931 The morphology of
Eupoterion pernix gen. nov., sp. nov. Uni. Calif. Publ. Zool.,
Vol. 36.
MiYASHiTA, Y. 1933 Drei neue parasitische Infusorien aus dem
Darme
einer Japanischen Siisswasseroligochaete. Annot. Zool.
Japon., Vol. 14.
Stevens, N. M. 1903 Further studies on the ciliate Infusoria,
Licnophora and Boveria. Arch. f. Protistenk., Vol. 3.
Chapter 37
Order 1 Holotricha Stein (continued)
630
HOLOTRICHA 631
Young trophont
Phoront
h,Spirophrya subpara-
Fig. 297. a, Foettingeria actiniarum, a trophont;
d, Syno-
sitica,a trophont, XlOOO; c, Phoretrophyra nebaliae, X1180;
phrya hypertrophica (Chatton and Lwoff).
Oospira
Genus Gymnodinioides Minkiewicz {PJujsophaga Percy;
Trophonts twisted along equatorial plane; gen-
Chatton and Lwoff).
erally 9 ciliary rows, in some a rudimentary row
between striae 5 and
6 at anterior end. Many species.
HOLOTRICHA 633
References
Chatton, E. and A. Lwoff 1935 Les cih^s apostomes. Arch. zool.
exp. et gen., Vol. 77.
Miyashita, Y. 1933 Studies on a freshwater foettingeriid ciliate,
Hyalospira caridinae n.g., n.sp. Japan. Jour. Zool., Vol. 4.
Chapter 38
Order 2 Spirotricha Butschli
With free cilia only; exceptionally with small groups of cirrus-like pro-
jections in addition to cilia
Uniformly ciliated; in Peritromidae dorsal surface without or with a
few cilia; in Licnophoridae cilia only on edge of attaching disk;
peristome usually extended; peristomal field mostly ciliated
Suborder 1 Heterotricha
Ciliation much reduced or none at all
Rounded in cross-section; cilia usually much reduced; adoral zone
encloses a non-ciliated peristomal field in spiral form
Suborder 2 Oligotricha (p. 652)
Compressed; carapaced; peristomal zone reduced to 8 membranellae
which lie in an oval hollow. .Suborder 3 Ctenostomata (p. 665)
Cirri only,on ventral side; dorsal side usually with rows of short bristles . .
636
SPIROTRICHA, HETEROTRICHA 637
*i..:^
3 4
it has been studied by several observers. Giese (1938) found that the
644 PROTOZOOLOGY
giants are carnivorous individuals feeding on small bacteria-fed in-
dividuals or other ciliates and remain large (with larger macronucle-
us and undulating membrane) after division, as long as appropriate
diet is supplied.
Genus Protocruzia Faria, da Cunha and Pinto. Peristome does
not turn right, leads directly into cytostome; convex left side not
ciliated, but bears bristles; flat right side with 3-5 faintly marked
ciliary rows; peristome begins at pointed anterior end and extends
1/4-1/3 the body length; cytopharynx (?); macronucleus simple;
contractile vacuole subterminal salt water. ;
P. pigerrima (Cohn) (Fig. 302, j). About 20/x (da Cunha); 50-
long (Kahl) peristome 1/4-1/3 the body length; salt water.
60^1 ;
in the colon of frogs and toads. Wichterman (1936) studied its life-
cycle in Hyla versicolor (p. 160).
1-2 mm. long; anterior end greatly expanded; the beautiful blue
color is due to a pigment, stentorin, lodged in interstriation gran-
ules; macronucleus beaded. Burnside (1929) studied its body and
nuclear sizes (p. 171).
S. striatus Barraud-Maskell. Dark bluish green; funnel-shaped;
peristomal edge irregularly undulating; striation conspicuous; macro-
nucleus beaded; up to 2.2 mm. long.
S. polymorphus (MiiWer) (Fig. 304,/). Colorless; with zoochlorellae;
1-2 mm. long when extended; macronucleus beaded; anterior end
expanded.
S. miilleri (Bory) (Fig. 304, g). Colorless; with zoochlorellae; 2-3
mm. long; anterior end expanded; posterior portion drawn out into
stalk, often housed in a gelatinous tube; on body surface 3-4 longer
and stiff cilia grouped among cilia; macronucleus moniliform.
*S. roeseli Ehrenberg (Fig. 304, h). 0.5-1 mm. long; anterior end
200m in diameter.
Genus Fabrea Henneguy. Pyriform; posterior end broadly round-
ed, anterior end bluntly pointed; peristome extends down from
anterior end 2/5 or more the body length, its posterior portion
closely wound; peculiar black spot beneath membranellae in anterior
portion of spiral adoral zone, composed of numerous pigment gran-
ules; without contractile vacuole; macronucleus, a sausage-shaped
body or in 4 parts; in salt water.
F. salina H. (Fig. 306, a, h). 120-220m by 67-125^ (Kirby); 130-
450/x by 70-200/x (Henneguy); cysts ovoidal, with gelatinous enve-
lope; 89-IIIm by 72-105^. Kirby (1934) found the organism in
ditches and pools in salt marshes, showing salinities 7.5-20.1 per
cent in California.
Genus Climacostomum Stein. Oval; flattened; right edge of peri-
stome without membrane, left edge, semicircular or spiral with a
strong adoral zone; peristomal field ciliated; cytopharynx a long
curved tube with a longitudinal row of cilia; macronucleus band-
form; contractile vacuole terminal, with two long canals; fresh or
brackish water.
C. virens (Ehrenberg) (Fig. 306, c). 100-300m long; with or without
P. arcHca D. (Fig. 306, /). Lorica about 430/i high, with spiral
ridge; off Norweigian coast 15-28 m. deep.
Genus ParafoUiculina Dons. Neck of lorica with a basal swelling;
attached either with posterior end or on a lateral surface; salt water.
References
Andrews, E. A. 1921 American Folliculinas : taxonomic notes.
Amer. Nat., Vol. 55.
1942 A folliculinid at Beaufort, North Carolina. Trans.
Amer. Micr. Soc, Vol. 61.
1944 FolliculiniH Protozoa on North American coasts.
Amer. Miclland Natural., Vol. 31.
Balamuth, W. Studies on the organization of ciliate Protozoa.
1941, Jour. Morph., Vol. 68; 1942, Jour. Exper. Zool., Vol. 91.
Hegner, R. 1934 Specificity in the genus Balantidium based on
size and shape of body and macronucleus, with description of
six new species. Amer. Jour. Hyg., Vol. 19.
Kahl, a. 1932 Urtiere oder Protozoa. In Dahl's Die Tierwell
Deutschlands. Part 15.
Kidder, G. W. 1937 The intestinal Protozoa of the wood-feeding
roach Panesthia. Parasitology, Vol. 29.
KiRBY Jr., H. 1934 Some ciliates from salt marshes in California.
Arch. f. Protistenk., Vol. 82.
Kudo, R. R. 1936 Studies on Nydotherus ovalis Leidy, with special
reference to its nuclear structure. Ibid., Vol. 87.
and P. A. Meglitsch 1938 On Balantidium praenucleatum
n.sp., inhabiting the colon of Blatta orientalis. Ibid., Vol. 91.
Levine, N. D. 1940 The effect of food intake upon the dimensions
of Balantidium from swine in culture. Amer. Jour. Hyg., Vol. 32.
McDonald, J. D. 1922 On Balantidium coli (Malmsten) and B.
suis (sp. nov.). Uni. Calif. Publ. Zool, Vol. 22.
Powers, P. B. A. 1935 Studies' on the ciliates of sea urchins. Papers
from Tortugas Lab., Vol. 29.
ScHMAHL, O. 1926 Die Neubildung des Peristoms bei den Teilung
von Bursaria truncatella. Arch. f. Protistenk., Vol. 54.
Stevens, N. M. 1903 Further studies on the ciliate Infusoria,
Licnophora and Boveria. Ibid., Vol. 3.
Strong, R. P. 1904 The clinical and pathological significance of
Balantidium coli. Bur. Gov. Labs. Manila, Biol. Lab., Bull. No.
26.
Chapter 39
Free-living
Oral portion of peristome lies free on ventral surface
Family 1 Halteriidae
Adoral zone encloses frontal peristomal field
Without lorica Family 2 Strobilidiidae (p. 653)
With lorica or test Family 3 Tintinnidae (p. 654)
Parasitic
Adoral and dorsal zones, both directed anteriorly and retractile; no
other cilia Family 4 Ophryoscolecidae (p. 654)
In addition to adoral and dorsal zones, groups of cirri in posterior half
of body, directed posteriorly and nonretractile
P'amily 5 Cycloposthiidae (p. 659)
652
;
near anterior end; posterior zone only half-spiral; in the caecum and
colon of horse. One species.
THE placed
ciliates
forms with a very sparse
group
in this are carapaced
ciHation. The
and compressed
adoral zone is also re-
duced to about 8 membranellae: These organisms are exclusively
free living and sapropelic in fresh, brackish, or salt water.
Posterior half of carapace with 4 ciliated rows on left and at least 2 rows
on right; with anterior row of cilia on left side. .Family 1 Epalcidae
Posterior half of carapace with cirrus-like groups on left only, none on
right; without frontal cilia
Long ciliated band extends over both broad sides
Family 2 Discomorphidae
Short ciliated band ventral, extending equally on both broad sides. . .
665
666 PROTOZOOLOGY
both lateral surfaces; 2 spines on right side; 2 cirrus-like groups on
posterior-left; sapropelic.A few species.
D. pectinata L. (Fig. 313, e,/). 70-90/x long; sapropelic.
References
Kahl, A. 1932 Ctenostomata (Lauterborn) n. subord. Arch. f.
Chapter 41
THE members
or are
cilia cirri
of this suborder are, as a rule, flattened
restricted to the ventral surface.
and strong
Except the fam-
ily Aspidiscidae, the dorsal surface possesses rows of short slightly
moveable tactile bristles. The peristome is very large with a well-
developed adoral zone. The cirri on the ventral surface are called,
according to their location, frontals, ventrals, marginals, anals
and caudals, as was mentioned before (Fig. 11, 6).
(transversals),
Asexual reporductionis by binary fission and sexual reproduction by
Family 1 Oxytrichidae
Ventrals and marginals not in longitudinal rows
Family 2 Euplotidae (p. 676)
No ventral cirri; caudal cirri Family 3 Paraeuplotidae (p. 677)
Adoral zone reduced Family 4 Aspidiscidae (p. 679)
668
SPIROTRICHA, HYPOTRICHA 669
tractile vacuole; salt water, with 5-20 per cent salt content. One
species.
C. koltzowii G. (Fig. 314, k, I). Band-form up to about 200/z long;
posteriorly attenuated forms up to about 100^ long.
Genus Psilotricha Oval to ellipsoid; frontals and anals un-
Stein.
and marginals long cirri, few; ventrals in 2
differentiated; ventrals
rows and a rudimentary row toward left; with or without zoochlo-
rellae; fresh water. A few species.
P. acuminata S. (Fig. 315, 6). 80-1 00)u long.
Genus Kahlia Horvath. Frontal margin with 3-4 strong cirri;
U. Umnetis Stokes (Fig. 315, d). About 200m long; fresh water
among vegetation.
U. longicaudatus S. (Fig. 315, e). About 200/x long; marsh water
with sphagnum.
U. dispar S. (Fig. 315,/). 150-170m long; fresh water.
a). About 160/i by 20m
U. halseyi Calkins (Fig. 317, peristome ;
b c ,((/////>. h
676 PROTOZOOLOGY
anals; marginals uninterrupted; 4-8 macronuclei; contractile vacu-
ole fresh water.
; One species.
0. grandis S. (Fig. 317,/). 100-300/x long.
Genus Onychodromopsis Stokes. Similar to Onychodromus ; but
flexible; 6 frontals of which the anterior three are the largest; fresh
water. One species.
O.fiexilis S. (Fig. 317, g). 90-125m long; standing pond water.
with 8 low ridges; peristome wide, with a small peristomal plate; end
of cytopharynx almost below the median ridge; 4th ridge between
anal cirri often extends to anterior end of body post-pharyngeal sac ;
a straight transverse line; 4th ridge between anals may reach anterior
end body; macronucleus C-shape with a flattened part in the left-
of
anterior region; micronucleus some distance from macronucleus at
anterior-left region; post-pharyngeal sac; fresh and brackish (salin-
ity 2.30 parts of salt per 1000) water.
E. plumipes Stokes. Similar to E. eurystomus. About 125/i long;
fresh water.
E. carinatus S. (Fig. 318, a). About 70/x by 50/i; fresh water.
E. Charon (Miiller) (Fig. 318, h). 70-90^ long; salt water.
Genus Euplotidium Noland. Cylindrical; 9 frontal-ventrals in 2
rows toward right; 5 anals; a groove extends backward from oral
region to ventral side, in which the left-most anal cirrus lies; peri-
stome opened widely at anterior end, but covered posteriorly by a
transparent, curved, flap-like membrane; adoral zone made up of
about 80 membranellae; longitudinal ridges (carinae), 3 dorsal and
2 lateral; a row of protrichocysts under each carina; a broad zone
of protrichocysts in antero-dorsal region; cytoplasm densely granu-
lated; salt water. One species.
E. agitatum N. (Fig. 318, c, d). 65-95^ long; erratic movement
rapid observed in half -dead sponges in Florida.
;
marked by a plate and with two ciliary tufts, near the middle of an-
terior half; 5-6 caudal cirri; macro nucleus curved band-form; a
terminal contractile vacuole; zooxanthellae, but no food vacuole in
cytoplasm; marine, on the coral.
P. tortugensis W. average indi-
(Fig. 319). Subcircular to ovoid;
viduals 85ju by 75m: with longer cilia; adoral
ciliary plate 37/1 long,
zone reaches nearly the posterior end; "micro nucleus not clearly
differentiated" (Wichterman) 5-6 caudal cirri about 13/i long; zoo-
;
681
682 PROTOZOOLOGY
water. Several species. Swarczewsky (1928) described several species
from Lake Baikal in Siberia.
S. gemmipara S. (Fig. 320, a). 80-120)u long; attached to the gill-
plates of Gammarus pulex and other species.
Genus Stylochona Kent. Peristomal funnel with an inner funnel.
One species.
S. coronata K. (Fig. 320, b). About 60^ long; on marine Gammar-
us.
Genus Kentrochona Rompel { Keritrochonopsis Doflein). Peri-
stomal funnel wide, simple, membranous; with or without a few (2)
spines.
K. nebaliae R. (Fig. 320, c). About 40ju long; much flattened, with
itsbroad side attached by means of gelatinous substance to epi-
and exo-podite of Nebalia geoffroyi; salt water.
Genus Heliochona Plate. Peristomal funnel with numerous needle-
like spines.
H. scheuteni (Stein) (Fig. 320, d). About 80-90^ long; on append-
ages of Gammarus locusta; salt water.
H. sessilis P. (Fig. 320, e). About 60/i long; on Gammarus locusta;
salt water.
Genus Chilodochona Wallengren. Peristome drawn out into two
lips; with a long stalk.
C. quennerstedti W. (Fig. 320, /). 60-1 15^* long; stalk, 40-160/x; on
Ebalia turnefacta and Portunus depurator; salt water.
Reference
Kahl, a. 1935 Uriiere oder Protozoa. In Dahl's Die Tierwelt
Deutschlands. Part 30.
Chapter 43
Order 4 Peritricha Stein
683
684 PROTOZOOLOGY
H. aesculacantha Jarocki and Jacubowska (Fig. 321, b). 30-52/i by
24-40/i in stagnant water.
;
PERITRICHA 685
of Cambarus.
Genus Rhabdostyla Kent. Similar to Epistylis; but solitary with
a non-contractile stalk attached to aquatic animals in fresh or salt
;
Fig. 323. a-c, Vorticella campamda (a, X400; b, part of stalk, X800;
c, telotroch, X200); d, e, V. convallaria (d, X400; e, X800); f-p, V. micro-
stoma {i, g, X400; h, X 840 i, telotroch, X400; j-p, telotroch-formation
;
PERITRICHA 691
692 PROTOZOOLOGY
L. vaginicola S. (Fig. 325, 1). Lorica 70^ by
48/x; attached to caudal
bristles and appendages minutus and Canthocamptus sp.
of Cyclops
L. patina Stokes (Fig. 325, w). Lorica 55/i by 50^; on Gammarus.
L. labiata S. (Fig. 325, n). Lorica 60/i by 55/i; on Gammarus.
THE Suctoria
Tentaculifera,
which have been also known as Acinetaria,
do not possess any cilia or any other cell-
etc.,
organs of locomotion in the mature stage. The cilia are present only
on young individuals which are capable of free-swimming, and lost
with the development of a stalk or attaching disk, and of tentacles.
Therefore, an adult suctorian is incapable of active movement. The
body may be spheroidal, elliptical, or dendritic and is covered with
;
fresh water.
T. salparum Entz (Fig. 327, c). On various tunicates such as
Molgula manhattensis; 40-60^ long; tentacles in 2 groups; salt water;
Woods Hole (Calkins).
T. columbiae Wailes (Fig. 327, d). 60-75m by 40-48/x in diameter;
cylindrical; tentacles at ends; nucleus spherical; in marine plankton;
Vancouver (Wailes).
T. micro-pteri Davis. Body elongate, irregular or rounded; up to
30-40m long by 10-12/i; fully extended tentacles 10-12;u long; cyto-
plasm often filled with yellow to orange spherules; a single micro-
nucleus; a single contractile vacuole; attached to the gill of small
mouth black bass, Micropterus dolomieu. Davis (1942) states that
when abundantly present, the suctorian may cause serious injury to
the host.
Genus Astrophrya Awerinzew. Stellate; central portion drawn out
into 8 elongate processes, each with a fascicle of tentacles; body cov-
ered by sand grains and other objects. One species.
A. arenaria A. (Fig. 327, e). 145-188^ in diameter; processes 80-
190ai long; in Volga river plankton.
Genus Lernaeophrya Perez. Body large; with numerous short pro-
longations, bearing very long multifasciculate tentacles; nucleus
branched; brackish water. One species.
L. capitata P. (Fig. 327, /). Attached to the hydrozoan, Cordy-
lophora lacustris in brackish water; 400-500^ long; tentacles 400^
long.
Genus Dendrosomides Collin. Branched body similar to Dendro-
soma, but with a peduncle; reproduction by budding of vermicular
form; salt water. One species.
D. paguri C. (Fig. 327, g). 200-300^ long; vermicular forms 350/i
long; on the crabs, Eupagurus excavatus and E. cuanensis.
SUCTORIA 697
706 PROTOZOOLOGY
stalk striated, about 150-270m long; tentacles in 4 groups; nucleus
ovoid; 1-3 contractile vacuoles; attached to the plastron of the tur-
tle.
wig); e, E. coronata, Xl40 (Kent); f, E. plana, front view, with two at-
tached Ophryocephalus, X35 (Wailes); g, Podocyathus diadema, X200
(Kent).
Collection
710
COLLECTION, CULTIVATION, OBSERVATION 711
Cultivation
A. Free-living Protozoa
work. Here only a few examples will be given. For further informa-
tion, the reader is referred to Belaf (1928), Needham et al. (1937),
etc.
Chromatophore-hearing fiagellates. —There are a number of culture
fluids. Two examples:
(a) Peptone or tryptone 2 .gm.
KH2PO4 0.25 gm.
MgS04 0.25 gm.
KCl 0.25 gm.
FeCla trace
Sodium acetate 2.0 gm.
Pyrex distilled water 1000 cc.
MgS04
714 PROTOZOOLOGY
If the culturewater becomes turbid, make subcultures or pour off
the water and fill with fresh distilled water or the solution. Culture
cium are added daily. Pace and Belda (1944) advocate the following
solution instead of distilled water:
K2HP04
COLLECTION, CULTIVATION, OBSERVATION 715
Magnesium sulphate
:
716 PROTOZOOLOGY
dium: 5 cc. of Bacto nutrient agar dissolved in boiling water, is
tubed and the tubes are autoclaved for 15 minutes at 15 pounds
pressure and slanted. After cooling, a modified Ringer's solution is
added to the agar slants to a height of 2-3 inches. The composition
of Ringer's solution used, is as follows
—
Trypanosoma and Leishmania. Novy, MacNeal and Nicolle
(NNN) medium: 14 gm. of agar and 6 gm. of NaCl are dissolved by
heating in 900 cc. of distilled water. When the mixture cools to about
50°C., 50-100 cc. of sterile defibrinated rabbit blood is gently added
and carefully mixed so as to prevent the formation of bubbles. The
blood agar is now distributed among sterile test tubes to the height
ofabout 3 cm., and the tubes are left slanted until the medium be-
comes solid. The tubes are then incubated at 37°C. for 24 hours to
determine sterility and further to hasten the formation of conden-
sation water (pH blood or splenic puncture containing
7.6). Sterile
Trypanosoma Leishmania is introduced by a sterile pipette
cruzi or
to the condensation water in which organisms multiply. Incubation
at 37°C. for trypanosomes and at 20-24°C. for Leishmania.
—
Entamoeba harreti. Barret and Smith (1924) used a mixture of
9 parts of 0.5% NaCl and 1 part of human blood serum. Incubation
at 10-15°C.
E. invadens. — Ratcliffe
and Geiman (1938) used a mixture of
gastric mucin "ground alum" salt 0.5 gm., and distilled
0.3 gm.,
water 100 cc. About 2 mg. of sterile rice starch is added to each cul-
ture tube at the time of inoculation. Culture at 20-30° C. and sub-
culture every 7 days.
E. histolytica and other amoebae of man. —The first successful cul-
COLLECTION, CULTIVATION, OBSERVATION 717
ture was made by Boeck and Drbohlav (1925) who used the follow-
ing media.
(a) Locke-egg-serum (LES) medium. The contents of 4 eggs
(washed and dipped in alcohol) are mixed with, and broken in, 50 cc.
of Locke's solution in a sterile flask with beads. The solution is made
up as follows:
NaCl 9 gm.
CaCl2 0.2 gm.
KCl 0.4 gm.
NaHCOa 0.2 gm.
Glucose •
2.5 gm.
Distilled water 1000 cc.
dium. Solid medium is the same as that of (a) or (b), but made up in
Ringer's solution which is composed of
NaCl 9 gm.
KCl 0.2 gm.
CaCla 0.2 gm.
Distilled water 1000 cc.
The medium is tubed, autoclaved, and slanted. The slants are cov-
ered with a 1 serum in 0.85% NaCl
:6 dilution of sterile fresh horse
solution. A 5 mm. loop of sterile rice flour or powdered unpolished
rice is added to each tube. In making subculture, remove 2 or 3 drops
of the rice flour debris from the bottom with a sterile pipette.
Plasmodium.— BsLSS and John's (1912) culture is as follows: 10 cc.
of defibrinated human blood containing Plasmodium and 0.1 cc. of
50% sterile dextrose solution are mixed in test tubes and incubated
at 37-39°C. In the culture, the organisms develop in the upper layer
of erythrocytes. Although several attempts have since been made
by several workers, Plasmodium has not been cultivated in vitro for
more than a few generations.
—
Balantidium coli. Barret and Yarb rough (1921) first cultivated
this ciliate in a medium consisting of 16 parts of 0.5% NaCl and 1
part of inactivated human
blood serum. The medium is tubed.
Inoculation of a small amount
of the faecal matter containing the
trophozoites is made into the bottom of the tubes. Incubation at
37°C. Maximum development is reached in 48-72 hours. Subcul-
tures are made every secondday. Reese used a mixture of 16 parts
of Ringer's solution and 1 part of Loeffler's dehydrated blood serum.
Atchley (1935) employed a medium composed of 4 parts of Ringer's
solution and 1 part of faeces, which is filtered after 24 hours, centri-
fuged and sterilized by passage through a Seitz filter. Nelson (1940)
also used 1 part of caecal contents of pig in 9 parts of Ringer's solu-
COLLECTION, CULTIVATION, OBSERVATION 719
Microscopical examination
A. Fresh preparations
ered by a depression slide with a thin coat of vaseline along the edge
of the depression, so as to make an air-tight compartment. In turning
over the whole, care must be taken to prevent the smaller circular
720 PROTOZOOLOGY
cover from touching any part of the sUde, as this would cause the
water to run down into the depression. Nemeczek (1926) seems to
have been the first one who used the second coverglass for this prepa-
ration. If the Protozoa to be examined are large and observation can
be made under a low power objective, the small coverglass should
be omitted.
As far as possible examine fresh preparations with low power ob-
jectives. The lower the magnification, the brighter and the larger the
field. The microscopical objects can quickly and easily be measured,
if an ocular micrometer division has been calculated in combination
with different objectives.
For observation of cilia, flagella, extruded polar filament of Micro-
sporidia, etc., the so-called changeable condenser is useful, since it
gives both bright and dark under dry objectives. The ordinary
fields
dark field condenser is used almost exclusively in conjunction with
an oil immersion objective and therefore for very active organisms
a great deal of time is often lost before satisfactory observation is
made.
When treated with highly diluted solutions of certain dyes, living
Protozoa exhibit some of their organellae or inclusions stained with-
out apparent injury to the organisms. These vital stains are usually
prepared in absolute alcohol solutions. A small amount is uniformly
applied to the slide and allowed to dry, before water containing Pro-
tozoa is placed on it. Congo red (1 1,000) is used as an indicator, as
:
its red color of the salt changes blue in weak acids. Janus Green B
(alkaline), cherry red (weak acid), and blue (strong acid). It also
stains nucleus slightly. Golgi bodies are studied in it, though its
specificity for this structure is not clear.
Parasitic Protozoa should be studied in the tissue or body fluids in
which they occur. When they are too small in amount to make a
suitable preparation, one of the following solutions may bq used.
Physiological salt solution. Widely used concentrations of NaCl
solutions are 0.5-0.7% for cold-blooded animals and 0.8-0.9% for
warm-blooded animals.
Ringer's solution. The one Dobell advocated has been given al-
cylinder, and let it stand for about 15 minutes. Remove the scum
floating on the surface and draw off the turbid fluid into another
cylinder, leaving the sediment and a little fluid just above it un-
touched. The majority of cysts are suspended in the drawn-off por-
tion of the emulsion. Centrifuge the fluid, pour off the supernatant
fluidand add water. Centrifuge again. Repeat this three times until
the supernatant fluid becomes clear. The sediment will be found to
contain more numerous cysts than small sample specimens.
B. Permanent preparations
a. Smear preparations
Smears are made either on coverglasses or slides. However, cover-
glass-smears are more properly fixed and require smaller amount of
reagents than slide-smears. Greater care must be excerised in han-
COLLECTION, CULTIVATION, OBSERVATION 723
O
%^
J Q) C3 %
Fig. 335. Sphaerita in a stained trophozoite of Entamoeba coli;
1,
The first two can be kept mixed without deterioration, but the acid
must be added just before fixation. Fix at room temperature or
724 PROTOZOOLOGY
warmed to 50°C. The fixative is placed in a square Petri dish and the
Bouin's fluid
Picric acid (saturated) 75 cc.
Formaldehyde 25 cc.
Fixation for 5-30 minutes; wash with 70% alcohol until picric acid
is completely washed away from the smears.
Sublimate-acetic
Saturated sublimate solution 100 cc.
This is the original fixative for Feulgen's nucleal reaction (p. 726).
Carney's fluid
Absolute alcohol 30 cc.
Osmium tetroxide
Fixation for 10-50 minutes; wash for one hour or longer in running
water.
The most commonly used stain is Heidenhain's iron haematoxy-
lin, as dependable and gives a clear nuclear picture, although it
it is
are decolorized in Petri dish in a diluted iron alum, 0.5% HCl in wa-
ter or 50% alcohol, or saturated aqueous solution of picric acid under
the microscope. Upon completion, the smears are washed thor-
oughly in running water for about 30 minutes. Rinse them in dis-
tilled water. Transfer them through ascending series of alcohol (50 to
The smears are now mounted in cedar wood oil (which is used for
immersion objectives) and the preparations should be placed in a
drying oven for a longer time than the balsam-mounted prepara-
tions.
Feulgen's nucleal reaction. The following solutions are needed.
(a) HCl solution. This is prepared by mixing 82.5 cc. of HCl (spe-
cific gravity 1.19) and 1000 cc. of distilled water.
Fuchsin-sodium bisulphite. Dissolve 1 gm. of powdered fuchsin
(b)
(basic fuchsin, diamant fuchsin or parafuchsin) in 200 cc. of distilled
water which has been brought to boiling point. After frequent shak-
ing for about 5 minutes, filter the solution when cooled down to 50°C.
into a bottle and add 20 cc. HCl solution. Cool the solution further
down to about 25°C. and add 1 gm. of anhydrous sodium bisulphite.
Apply stopper tightly. Decolorization of the solution will be com-
pleted in a few hours, but keep the bottle in a dark place for at least
24 hours before using it.
(c) Sulphurous water.
FiG. 336. Diagrams showing how a thin blood film is made on a slide.
cated. For this, 4 to 6 drops of blood are placed in the central half-
inch square area, and spread them into an even layer with a needle
or with a corner of a slide. Let the film dry. With a little practice, a
satisfactory thick smear can be made. It will take two hours or more
to dry. Do not dry by heat, but placing it in an incubator at 37°C.
will hasten the drying. When thoroughly dry, immerse it in water
and dehaemoglobinize it. Air dry again.
Thin and thick film. Often it is time-saving if thin and thick films
are made on a single slide. Place a single drop of blood near the center
COLLECTION, CULTIVATION, OBSERVATION 729
and make a thin film of it toward one end of the sUde. Make a small
thick smear in the center of the other half of the slide. Dry. When
thoroughly dry, immerse the thick film part in distilled water and de-
haemoglobinize it. Let the slide dry.
Blood smears must be stained as soon as possible to insure a proper
staining, as lapse of time or summer heat will often cause poor stain-
ing especially of thick films. Of several blood stains, Giemsa's and
Wright's stains are used here. For staining with Giemsa's stain, the
thin film is fixed in absolute methyl alcohol for 5 minutes. Rinse well
the slide in neutral distilled water. After shaking the stock bottle
(obtained from reliable makers) well, dilute it with neutral distilled
water in a ratio of one drop of stain to 1-2 cc. of water. Mix the solu-
tion and the blood film is placed in it for 0.5-2 hours or longer if
c. Section preparations
731
—
732 PROTOZOOLOGY
Aggregatidae, 465, 466-469 Amoeba continued
Aging in Protozoa, 166-170 limicola, 346, 347
Agriodrilus, 559 meleagridis, 266
Agriolimax agrestis, 30, 601 mira, 104
Agrion puella, 447 proteus, 24, 42, 44, 52, 69, 70, 72,
Ahlstrom, 228 87, 88, 90, 92, 94, 103, 105, 107,
Aikinetocystidae, 431, 434 108, 109, 110, 141, 142, 143, 170,
Aikinetocystis, 434 345,34^,713
singularis, 434, 435 radiosa, 18, 42, 87, 347, 348
Akaryomastigont, 315 spumosa, 347-348
Alasmidonta undulata, 624 striata, 39, 42, 73, 345-346
Albertisella, 436 verrucosa, 18, 21, 22, 39, 103, 105,
crater, 436 106, 107, 170, 345, 346
Alexeieff, 130 vespertilio, 347, 348
Algae, 91, 92, 329, 330, 331, 333, Amoebiasis, 355, 357, 365
709 Amoebic dysentery, 354, 357, 364
Alisma, 342 Amoebidae, 343, 344, 345-351
AUantocystidae, 431, 440 Amoebina, 195, 329, 343-371
Allantocystis, 440 Amoebodiastase, 91
dasyhelei, 439, 440 Amoeboid movements, 106-110, 114,
Allantosoma, 705 328
brevicorniger, 705, 707 Amphacanthus, 655
dicorniger, 705, 707 ovum-rajae, 655, 656
intestinalis, 705, 707 Amphibia, 25, 239, 267, 279, 299, 301,
AUegre, 236 313, 365, 367, 473, 475, 476, 480,
Allman, 65 505, 521, 527, 547, 548, 549, 550,
AUogromia, 374 555, 685, 692
AUoiozona, 578 Amphidinium, 253
trizona, 578 fusiforme, 253
Allolobophora caliginosa, 557 lacustre, 21, 253, 253
AUomorphina, 404 scissum, 252, 253
trigona, 404 Amphileptidae, 580-582
AUosphaerium, 590 Amphileptus, 23, 65, 580
caudatum, 590 branchiarum, 580, 581
convexa, 127, 590 claparedei, 21, 580, 581
granulosum, 590 meleagris, 580
palustris, 589, 590 Amphilonche, 421
sulcatum, 590 hydrometrica, 421
Allurus tetraedurus, 557 Amphilonchidae, 420
Aloricata, 683-689 Amphimonadidae, 268, 285-287
Alveolinella, 400 Amphimonas, 285
mello, 399 globosa, 285, 286
Alveolinellidae, 400 Amphimixis, 161
Amara axigustata, 452 Amphionts, 436
Amaroucium, 459 Amphioxus, 608
Amaurochaete, 339 Amphipoda, 630
fuliginosa, 339 Amphisiella, 670
Amaurochaetidae, 339 thiophaga, 669, 670
Amberson, 102 Amphisteginidae, 403
Ambystoma tigrinum, 549 Amphitrema, 388
Ameuirus albidus, 568 flavum, 387, 388-389
Amiba, 345 Amphiura squamata, 634
Amitosis, 122-130 Amphizonella, 382
Ammodiscidae, 398 violacea, 382
Ammodiscus, 398 Amphorocephalus, 454
incertus, 397, 398 amphorellus, 453, 454
Amoeba, 6, 18, 19, 90, 91, 114, 115, Amphoroides, 452
116, 117, 176,345,713 calverti, 450, 452
discoides, 105, 345, 346 AmpuUacula, 578
dofleini, 87 ampulla, 578
dubia, 44, 87, 90, 91, 105, 345, 346 Anabolic products, 98-101
gorgonia, 347, 348 Anacharis, 701
guttula, 42, 346 Anal cirri, 49, 50, 51
AUTHOR AND SUBJECT INDEX 733
This Book
(3rd Edition)
PROTOZOOLOGY
By Richard R. Kudo, D.Sc.