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chambers of the Foraminifera, the siliceous and very ornamental framework of the
Radiolaria, and the chitinous coat of many Flagellata, Infusoria, etc. Some forms
make use of foreign bodies found in their surroundings, such as grains of sand, to
construct their protective coverings.
The food often consists of small animal or vegetable organisms and of organic
waste; it is usually introduced in toto into the endoplasm. On the other hand, the
Suctoria extract nourishment from their prey by means of their tentacles. Many
parasitic species also ingest solid food, others feed by endosmosis.
In all cases one nucleus at least is present. It is true that the existence of non-
nucleated Protozoa, the so-called Monera, is still insisted upon, but some of these
have already proved to be nucleated, and the presence of nuclei in the others will
no doubt be established. Very often the number of nuclei increases considerably,
but these multinucleate stages are always preceded by uninucleate stages. In the
Infusoria, in addition to the larger or principal nucleus (macronucleus) there is
usually a smaller reproductive nucleus (micronucleus). This dualism of the nuclear
apparatus is considered by some to be general, and usually to appear first at the
onset of reproduction.
The form and structure of the nucleus vary greatly in different species. There are
elongate, kidney-shaped, or even branched nuclei as well as spherical or oval
ones. In addition to vesicular nuclei with a distinct karyosome and incidentally also
with a nuclear membrane, homogeneous and more solid formations are frequently
encountered. The nuclei are always differentiated from the protoplasm by their
reactions, particularly in regard to certain stains.
In many Protozoa an extra-nuclear mass, sometimes compact, sometimes diffuse,
arises from or near the nucleus. This mass, whose staining reactions resemble
those of the nucleus, is termed the chromidial apparatus. On the dualistic
hypothesis, two varieties of chromidia occur, one originating from the vegetative
nucleus (macronucleus), being chromidia in the restricted sense, the other derived
from the reproductive or micronucleus being termed sporetia. Chromidia consist of
altered (? katabolic) nuclear material.
The nucleus plays the same part in the life of the single celled organisms as it
does in the cells of the Metazoa and Metaphyta. It appears to influence in a
certain manner all, or at least most, of the processes of life, such as motility,
regeneration, growth, and generally also digestion. Its principal influence,
however, is exercised in the propagation of the cells, as this is always brought
about by the nucleus.
The propagation of the Protozoa is effected either by division or by means of direct
budding. In division, which is preceded by direct or indirect (mitotic) division of
the nucleus, the body separates into two, several, or even a great many segments.
In this process the entire substance of the body is involved, or a small residual
fragment may be left, which does not undergo further division and finally perishes.
In the budding method of multiplication a large number of buds are formed, either
on the surface or in the interior of the organism. Where divisions or buddings
follow one another rapidly, without the segments separating immediately after
their production, numerous forms develop, which are often unlike the parental
forms, and these are termed swarm spores or spores. Divisions imperfectly
accomplished lead to the formation of protozoal colonies.
Sometimes encystment 9 takes place previous to division. Frequently, also, sexual
processes appear, such as the union of two similar (isogamous) or dissimilar
(anisogamous) individuals. In the latter case sexual dimorphism occurs, with the
formation of males (microgametes) and of females (macrogametes). The union
may be permanent (copulation), the process being comparable with the
fertilisation of the ovum by a spermatozoon. On the other hand, attachment may
be transient (conjugation) when, after the exchange of portions of the nucleus,
the couple separate, to multiply independently of each other. Sometimes there is
an alternation of generations, as there may be several methods of propagation
combined in the same species, either direct multiplication, conjugation, or
copulation being practised; the different generations may thus, in certain cases, be
unlike morphologically.
Protozoa inhabit salt water as well as fresh water; they are also found on land in
very damp places, and invade animals as parasites.

Classification of the Protozoa.

Class I.—Sarcodina (Rhizopoda). Protozoa, the body substance of which forms
pseudopodia; many of them are capable of developing chitinous, chalky, or
siliceous coverings or skeletal structures, which, however, permit the protrusion of
the pseudopodia either over the entire periphery or at certain points. They possess
one nucleus or several.
Order 1.—Amœbina (Lobosa) naked or with a simple shell, sometimes formed
of a foreign substance; the pseudopodia may be lobose or finger-
shaped; there may be a contractile vacuole; generally only one
nucleus. They live in fresh or salt water, in the soil, and also
parasitically.
 Order 2.—Foraminifera (Reticularia). Mostly provided with a calcareous shell,
usually consisting of several chambers, and allowing the protrusion of
the pseudopodia either at the periphery or only at the opening. The
pseudopodia are filamentous and frequently anastomosed; there is no
contractile vacuole; there are usually several nuclei. Mostly marine.
Order 3.—Heliozoa. Naked, or with a chitinous or simple radial siliceous
skeleton; the pseudopodia are filamentous, and are frequently
supported by firmer axes, which exhibit no tendency to anastomosis;
there is a contractile vacuole; one or several nuclei. Live in fresh water.
Order 4.—Radiolaria. The body has radially-disposed filamentous
pseudopodia, and the nucleus is hidden in the central capsule; there is
almost always a siliceous framework, consisting of pieces arranged
radially, tangentially, or lattice-like; there is no contractile vacuole, but
fluid-containing hydrostatic vacuoles are present in the peripheral
protoplasm. Marine.
Class II.—Mastigophora (Flagellata). Protozoa with one or several long flagella
used for locomotion and for acquiring food; in stationary forms their only function
is to take in food. Cytostome and contractile vacuole may be present. May be
either naked or provided with protective coverings; one or more nuclei. They live
either in fresh or salt water, or may be parasitic.
This class is again divided into several sub-classes and orders, of which only the
Euflagellata, with the Protomonadina and Polymastigoda are of interest here.
Class III.—Sporozoa. Protozoa that only live parasitically in the cells, tissues, or
organs of other animals. They ingest liquid food by osmosis; the surface of the
body is covered with an ectoplasmic layer, or cuticle; they have no cilia in the adult
state, but may form pseudopodia. Flagella occur, but only on the male propagating
individuals. There may be one or numerous nuclei, but no contractile vacuole.
Propagation by means of spores, mostly provided with sporocysts, is characteristic.
Sub-class 1.—Telosporidia. These are usually of constant form, rarely
amœboid; they are uninucleate in the mature state; they live within
host cells in the first stage. Spore-formation occurs at the end of the
life-cycle.
Order 1.—Gregarinida. Body of a constant, usually elongate form, surrounded
by a cuticle. In the early stage they lead an intracellular existence; in
the mature stage they live within the intestine or body cavity of
invertebrate animals, especially the Arthropoda, and, like intestinal
parasites, are provided with clinging organs. Copulation usually
 isogamous; the spores have coats (chlamydospores) and usually
contain several minute germs (sporozoites).
Order 2.—Coccidiidea. Body of uniform spherical or oval shape: they lead an
intracellular life, but are not freely motile in cavities of the body.
Fertilization is anisogamous; the spores have coats or shells
(sporocysts), and usually contain several sporozoites. Exhibit
alternation of generations.
Order 3.—Hæmosporidia. Parasites of the blood corpuscles of vertebrate
animals; they exhibit amœboid movement; fertilization is anisogamous;
many present alternation of generations and hosts; spores naked.
Sub-class 2.—Neosporidia. They are multinucleate when adult, and the form
of the body varies exceedingly (often amœboid); spore-formation
commences before the completion of growth.
Order 1.—Myxosporidia. The spores have valvular coats, with or without
caudal appendages, with two, rarely four, polar capsules. They live free
in such organs as the gall or urinary bladder, but are chiefly found in
connective tissue. They occur especially in fishes.
Order 2.—Microsporidia. Spores with coats or sporocysts; no caudal
appendage, with one polar capsule. They usually live in the tissues of
Arthropoda.
Order 3.—Sarcosporidia. Elongate parasites of the muscular fibres of amniotic
vertebrates, on rare occasions they occur also in the connective tissue;
the spores, which are kidney or sickle-shaped, are naked and
apparently have no obvious polar capsule.
Order 4.—Haplosporidia. Simple organisms, forming simple spores; they occur
in Rotifers, Polychætes, Fish and Man.
Class IV.—Infusoria (Ciliata). The body is generally uniform in shape, with cilia
and contractile vacuole, frequently also with cytostome; usually has macro- and
micro-nucleus; live free in water and also parasitically.
The orders Holotricha, Heterotricha, Oligotricha, Hypotricha and Peritricha are
classified according to the arrangement of the cilia.
Class V.—Suctoria. Bodies with suctorial tubes, contractile vacuoles, macro- and
micro-nucleus, no cytostome. They generally invade aquatic animals as cavity
parasites, yet also attack plants; early stage ciliated. Live sometimes as parasites
on Infusoria. [The Suctoria are frequently regarded as a sub-class of the
Infusoria.]
The Protozoa and Protophyta are sometimes united under the term Protista
(Haeckel, 1866). The Spirochætes are Protists (see pp. 114–128).

Class I. SARCODINA, Bütschli, 1882.

Order. Amœbina, Ehrenberg.
A. Human Intestinal Amœbæ.

The first record of the occurrence of amœba-like organisms in the human

intestine, that is, in intestinal evacuations, was that of Lambl (1859); nevertheless,
the case was not quite conclusive, as the occurrence of testaceous amœbæ of
fresh water (Arcella, Difflugia) was also reported. In 1870 Lewis found amœbæ
associated with disorders of the large intestine in patients in Calcutta. A year later
Cunningham reported from the same locality that he had observed on eighteen
occasions, in one hundred examinations of dejecta from cholera patients,
colourless bodies with amœboid movements, which became encysted and
multiplied by fission. The daughter forms were said to be capable of dividing
again, but they might also remain in contact. Contractile vacuoles were not
noticed. The same bodies were observed also in simple diarrhœa (twenty-eight
cases out of one hundred.)
The case reported by Lösch in 1875 attracted
more attention. It was that of a peasant,
aged 24, who came from the province of
Archangel. He was admitted into Eichwald’s
clinic at Petrograd with symptoms of
dysentery. In the discharges containing blood
and pus, Lösch found amœbæ in large
numbers. When at rest these amœbæ
measured from 20 µ to 35 µ; in a state of
movement their length might extend up to Fig. 1.—Amœba coli, Lösch, in
60 µ (fig. 1). The pseudopodia appeared only the intestinal mucus. (After
singly, and, since they were hyaline Lösch.)
(ectoplasmic), were thus distinguished from
the markedly granular endoplasm that enclosed a spherical nucleus of from 5 µ to
7 µ in diameter. One or more non-contractile vacuoles were present. Quinine
enemata had the effect of making the amœbæ disappear from the fæces and thus
causing the diarrhœa to abate. Four months after admission the patient died from
the results of intercurrent pneumonia. At the autopsy ulceration of the large
intestine was found, especially in the lower parts. Lösch connected the amœbæ
with the ulcerations by experiments made on four dogs by injecting them with
recently passed stools (per os et anum). Eight days after the last injection
numerous amœbæ were found in the fæces of one of these dogs; eighteen days
after the injection the animal was killed. The mucosa of the rectum was inflamed,
covered with blood-stained mucus and ulcerated in three places. Numbers of
amœbæ were found both in the pus of the ulcers and in the mucus. The three
other dogs remained healthy. From these observations Lösch concluded that the
species of amœba described by him as Amœba coli could not be regarded as the
primary cause of the disease, but that it was certainly capable of increasing a
lesion of the large intestine already present, or at least of preventing its healing.
B. Grassi (1879) found in the stools of healthy as well as in those of diarrhœic
patients from various localities in Northern Italy, amœbæ similar to those
discovered by Lösch. As this was of frequent occurrence, the pathogenicity could
not be definitely established. Normand, formerly naval surgeon at Hong-Kong,
observed numerous amœbæ in the dejecta of two patients suffering from colitis.
Many further investigations, which cannot be quoted in detail, showed not only
that intestinal amœbæ were widely distributed in man, but indicated with greater
certainty their rôle as agents of dysentery. The Commission sent out by the
German Government in the year 1883 to investigate cholera in India and Egypt—
whose members discovered the cholera bacillus—also collected information with
regard to dysentery. In five cases of dysentery examined post mortem at
Alexandria, with the exception of one case in which ulceration of the colon had
already cicatrized or was approaching cicatrization, R. Koch found amœbæ as well
as bacteria in sections from the base of the ulcers, although such had previously
escaped notice in examination of the dejecta. Encouraged by these results,
Kartulis (1885), who had discovered amœba-like bodies in the stools of patients
suffering from intestinal complaints at Alexandria, continued his investigations.
The results, obtained from more than 500 cases, gave rise to the theory that
typical dysentery was caused by amœbæ as were also the liver-abscesses that
often accompany it. Kartulis supported his theory not only by the regular
occurrence of amœbæ in the stools of dysenteric patients and their absence in
other diseases, and by the occurrence of the parasites in ulcers of the large
intestine and in the pus from liver-abscesses, but also by experiments which he
performed on cats. These were infected by injection per anum of stool material
rich in amœbæ from subjects of dysentery. The infection took place also when
amœba-containing, but bacteria-free, pus from liver-abscesses was used. It has
been objected that the infection of man with Amœba coli, as the dysenteric
amœbæ were then generally designated, does not take place per anum but per
os. This difficulty, however, diminished in proportion as the encysted states of
amœbæ (fig. 2), long known in the case of other Protozoa, became understood.
The infection of man (Calandruccio, 1890) and of cats (Quincke and Roos)
succeeded solely when material containing such stages was used. Amœbæ
introduced into the intestine multiply there by fission (Harris, 1894). However, this
theory, to which various other authors gave support on the grounds of their own
observations, encountered opposition. Thus it was established that amœbæ were
not found in patients in every place where dysentery was endemic, or else they
were much rarer than was expected. Further, amœbæ were present in the most
varied kinds of intestinal diseases, both of infective and non-infective characters.
Also they were present in quite healthy persons.
Moreover, for various reasons, infection experiments on animals failed to supply
proof, and finally a bacterium was discovered (Shiga, 1898) to be the excitant of
one form of dysentery. Agglutination attested the specific part played by this
organism, as it was produced by the blood serum of a person suffering from or
recovered from dysentery, but not by the serum of one who was uninfected.
Bacillary dysentery consequently was a distinct entity. The final step to be taken
was to decide whether there was a specific amœbic enteritis (amœbic dysentery
or amœbiasis, according to Musgrave).
This question should decidedly be regarded from
the positive point of view. It is intimately connected
with another, namely, whether there are not several
species of intestinal amœbæ. The possibility of this Fig. 2.—Encysted intestinal
had already been recognized. In addition to the amœbæ show­ing nuclear
Amœba coli Lösch, R. Blanchard distinguished yet multi­pli­ca­tion. (After B.
another, Amœba intestinalis, and designated Grassi.)
thereby the large amœbæ described in the first
communication made by Kartulis; later on he stated the distinction between the
species. Councilman and Lafleur 10 (1891) considered the amœba of dysentery to
be Amœba coli Lösch and so re-named the species Amœba dysenteriæ. Kruse and
Pasquale (1893) employed the same nomenclature, but retained the old name
Amœba coli Lösch for the non-infectious species. Quincke and Roos (1893) set
forth three species: a smaller species (25 µ) finely granular, pathogenic for men
and cats (Amœba coli Lösch); a larger species (40 µ) coarsely granular, pathogenic
for men but not for cats (A. coli mitis); and a similar species non-pathogenic either
for man or cat (A. intestini vulgaris). Celli and Fiocca (1894–6) went still further,
they distinguished:
(1) Amœba lobosa variety guttula (= A. guttula Duj), variety oblonga (= A.
oblonga Schm.) and variety coli (= A. coli Lösch).
(2) Amœba spinosa n. sp. occurring in the vagina as well as in the intestine of
human patients suffering from diarrhœa and dysentery.
(3) Amœba diaphana n. sp. found in the human intestine in cases of dysentery.
(4) Amœba vermicularis Weisse, present in the vagina and in dysentery; and
(5) Amœba reticularis n. sp. in dysentery.
Shiga distinguished two species; a larger pathogenic species with a somewhat
active movement, and a small harmless species with a somewhat sluggish
movement. Bowman mentions two varieties, Strong and Musgrave (1900) two
species—the pathogenic Amœba dysenteriæ and the non-pathogenic Amœba coli;
Jäger (1902) and Jürgens (1902) mention at least two species. In the following
year (1903) a work by Schaudinn was published which marked a real advance.
This, in conjunction with the establishing of a special genus (Endamœba or
Entamœba) for human intestinal amœbæ first by Leidy 11 and then by Casagrandi
and Barbagallo, 12 for the time cleared up the confused nomenclature, the old
name Amœba coli being retained for the harmless intestinal amœbæ of man,
whereas the pathogenic species was designated Entamœba histolytica. The history
of more recent work is incorporated in the accounts of the entamœbæ given
below.

Entamœba coli, Lösch, 1875, emend. Schaudinn, 1903.

Syn.: Amœba coli, Lösch, 1875. Entamœba hominis, Casagr. et Barbag. 1897.

The amœboid trophozoite, according to Lösch, measures 26 µ to

30 µ and upwards; according to Grassi 8 µ to 22 µ; according to
Schuberg 12 µ to 26 µ. A separation of the body substance into
ectoplasm and endoplasm is only perceived during movement. The
pseudopodia, which are generally only protruded singly, are broad
and rounded at the end (lobopodia) and are hyaline, while the
remainder of the body is granular. The ectoplasm is less refractile
than the rest of the cytoplasm; it also stains less intensely (fig. 1),
and is best seen on protrusion of a pseudopodium. Red blood
corpuscles are rarely, if ever, found ingested in the cytoplasm.
The nucleus is vesicular, and is
spherical when inactive, measuring
5 µ to 7 µ, with a thick nuclear
membrane. In the centre of the
nucleus is a chromatinic body or
karyosome or sometimes several
small nuclear bodies formed of plastin
and chromatin; the remaining
chromatin is arranged on the
achromatic network in the form of
Fig. 3.—Entamœba coli: life-
fine granules, especially thickly cycle, a-e, stages in binary
deposited on the nuclear membrane. fission; A-D, schizogony, with
formation of eight merozoites;
Entamœba coli lives as a commensal 2–10, cyst formation or
in the upper portion of the large sporogony, with formation of
intestine, where the fæces still eight nucleate cysts. (After
possess a pulpy consistency. With Castellani and Chalmers)
their concentration and change in
reaction lower in the bowel, the parasites either die or else if they
are at a suitable stage of development form resistant cysts. These
cysts (fig. 2) can be found in great abundance in normal fæces, as
Grassi first observed. Slight laxantia or intestinal diseases of any kind
producing increased peristalsis, however, show amœbæ even in the
unencysted condition, provided that the person harbours intestinal
amœbæ generally. The intensity of infection varies according to the
locality; thus Schaudinn found that 50 per cent. of the persons
examined were infected with harmless amœbæ in East Prussia, 20
per cent. in Berlin and about 66 per cent. on the Austrian littoral.
The life-history (fig. 3) of the parasite exhibits two phases: (a)
asexual multiplication in the intestine, either by binary fission or by
schizogony with formation of eight merozoites, and (b) sporogony
leading to the production of eight-nucleate cysts. Infection results
from ingestion of cysts. Only cysts with eight nuclei are infective.
The diameter of such cysts is about 15 µ to 20 µ.
There are varying accounts of the details of the life-cycle of Entamœba coli in its
different stages. Thus, regarding schizogony or multiple fission it was formerly
stated that the nucleus of the parent amœba divided into eight portions, which
after dissolution of the nuclear membrane, passed outwards into the cytoplasm,
which segregated around each. Eight merozoites were thus produced. More
recently the process of schizogony has been considered to consist in the repeated
division of the nucleus into two, four, and finally eight nuclei (fig. 3, a-d), and the
formation of eight merozoites or amœbulæ.
The process of encystment is initiated by the extrusion of all liquid and foreign
bodies from the protoplasm, which assumes a spherical form (fig. 4, a). The
rounded uninucleate amœba then secretes a soft gelatinous coat, which finally
differentiates into a double contoured cyst wall in older cysts. According to
Casagrandi and Barbagallo, the size of the cyst varies from 8 µ to 30 µ, and
averages about 15 µ. According to Schaudinn (1903) the cytological changes
during cyst formation are as follows. The nucleus of a rounded uninucleate form
divides into two (fig. 4, b). Each of these nuclei fragments into chromidia (fig. 4,
c), some of which are absorbed, while others reunite so that the cell becomes
binucleate again. Each of these nuclei, by a twice repeated division, produces
three nuclei (fig. 4, d), the smaller two of which degenerate and were regarded as
reduction nuclei. There is a clear zone or vacuole in the middle of the cyst during
these maturation processes, dividing the cyst into two halves. After the nuclear
reduction the clear space disappears, and each nucleus (termed by some a
gamete nucleus) divides into two pronuclei (fig. 4, e). The pronuclei of the pairs
were said by Schaudinn to differ slightly. Copulation occurs between pairs of unlike
pronuclei, and is an example of autogamy (fig. 4, f). When complete, each of the
fusion nuclei (synkarya) divides twice, giving rise first to four and finally to eight
nuclei. Eight amœbulæ are thus formed within the cyst.
According to Hartmann and Whitmore (1911) 13, however, autogamy does not
occur within the cysts of E. coli. They consider that eight small amœbulæ are
formed (fig. 3, 2-10) which escape from the cyst and then conjugate in pairs
(fig. 3, 10-12), afterwards growing into a new generation of trophozoites.
Only some 10 to 20 per cent. of the cysts evacuated with the fæces undergo the
full course of development, the majority perish previously. In old dry fæces, only
cysts with eight nuclei are found, and it is these alone that cause the infection.
Entamœba williamsi, E. bütschlii, E. hartmanni and E. poleki (Prowazek) are
probably only varieties of E. coli.

The principal feature distinguishing

Entamœba coli from E. histolytica is
the formation of eight-nucleate cysts
by the former as contrasted with the
tetra-nucleate cysts of the latter. The
cyst-wall of E. coli is thicker than that
of E. histolytica (tetragena). Further,
E. coli does not usually ingest red Fig. 4.—So-called autogamy of
blood corpuscles, nor are “chromidial Entamœba coli. a, rounded
amœba; b, nucleus dividing; c,
blocks” present inside its cyst (see
the two daughter-nuclei giving
p. 40). off chromidia; d, each nucleus
According to Chatton and Lalung- has formed two reduction nuclei;
e, cyst membrane formed, and
Bonnaire 14 (1912) the entamœbæ of
gamete nuclei are dividing; f,
vertebrates should be placed in a cyst with two synkarya.
separate genus Löschia, as they differ
in their life-history from E. blattæ, the type species of Entamœba.
Leidy (1879), however, named the genus Endamœba, but further
researches are necessary on biological variation among these
organisms.

Entamœba histolytica, Schaudinn, 1903.

Syn.: Amœba coli, autt. p. p. Amœba dysenteriæ, autt. p. p.

The average size of the amœboid trophozoite is 25 µ to 30 µ. In

fæces diluted with salt solution the amœbæ swell to 40 µ and more.
There is sometimes separation of the body substance into a strongly
refractile vitreous ectoplasm and a corneous endoplasm, pronounced
even in repose, although the former is not equally thick at all parts
of the periphery. In the endoplasm generally there are numerous
foreign bodies (bacteria, epithelial cells, colourless and red blood
corpuscles (fig. 6), and occasionally living flagellates of the
intestine). The nucleus is 4 µ to 6 µ in diameter, and may be difficult
to recognize because it is sometimes weakly refractile and poor in
chromatin. Its shape is slightly variable; it is usually excentric,
sometimes wholly peripheral at the limit of the two parts of the
body. Vacuoles are not present in quite fresh specimens, but appear
later. In the study of E. histolytica, the morphological characters of
the trophozoite or vegetative stage of the organism formerly
separated as E. tetragena (figs. 5, 6, 8a) must be considered (see
p. 38).

Fig. 5.—Entamœba histolytica (tetragena form), showing three

successive changes of form due to movement. × 1100. (After
Hartmann.)

The history of the development of these species, which give rise to amœbic
enteritis as distinguished from bacillary dysentery, was formerly not so well known
as that of E. coli. Upon being introduced into cats (per anum) dysenteric amœbæ
provoke symptoms similar to those in man. In the latter, besides metastatic liver
abscesses, abscesses of the lungs, and, according to Kartulis, cerebral abscesses
are occasionally produced. Marchoux (1899) states that when the disease has
lasted for some time liver abscesses are produced in cats also.
 Fig. 6.—Entamœba histolytica which has ingested
many red blood corpuscles. × 1100. (After
Hartmann.)

Fig. 7.—Section through wall of large intestine (of a man) close

under an ulcer caused by Entamœba histolytica. A, amœbæ that
have penetrated partly in blood-vessels (Bv), partly in tissue of
submucosa to the muscularis. Magnified. (After Harris.)
In the large intestine of infected cats the amœbæ creep over the epithelium, and
here and there they force the epithelial cells apart, as well as removing them or
pushing them in front of them; the amœbæ thus insert themselves into the
narrowest fissures. They penetrate also into the glands through the epithelium,
and thence into the connective tissue of the mucosa. Intestinal and glandular
epithelia perish under the influence of these parasites: the cells are pushed aside,
fall to pieces or are absorbed by the amœbæ. In the connective tissue of the
mucosa the amœbæ migrate further, and often accumulate above the muscles.
Finally they rupture this and force their way into the submucosa. In cats,
apparently, the penetration is not so great as in men, according to Kruse and
Pasquale. During their migration the parasites also gain access to the lymph-
follicles of the wall of the intestine, which become swollen and commence to
suppurate; follicular abscesses arise and after their rupture follicular ulcers. The
diseased patches in the mucosa are markedly hyperæmic and numerous
hæmorrhages are set up. Roos and Harris state that the amœbæ also penetrate
into the blood-vessels (fig. 7) and this explains the occurrence of metastatic
abscesses. 15 The whole submucosa is severely swollen at the diseased spot and
undergoes small-celled infiltration in the neighbourhood of the colonies of
amœbæ. From these findings Jürgens (1902) draws the conclusion 16 which is
followed here, that the amœbæ are causative agents of the enteritis of cats, which
disease is well defined, both pathologically and anatomically. Subsequent
researches confirm the experience of earlier authors; great precautions were taken
to exclude errors, hence, as with Gross and Harris, no exception can be taken to
their results. The inoculation material was derived from soldiers who suffered from
amœbic enteritis in China and who were admitted into the garrison hospital at
Berlin. In order to be independent of the patients themselves, transmission
experiments from cat to cat were performed, after the first experiments on cats
yielded positive results. This was also effected by rectal feeding as employed by
earlier workers. Such appeared necessary in order to prevent the evacuation of the
inoculation material per anum, as well as to avoid the employment of morphia and
ether narcosis. Forty-six cats were used for the experiments. Ten cats received
tested stools containing motile amœbæ from soldiers suffering from amœbic
enteritis contracted in China. Sixteen other cats received stools from cats infected
by inoculation. All the animals sickened and suffered from the disease. Five cats
received dejecta from human amœbic enteritis in which, however, no motile
amœbæ were present. Thirteen cats received stools from soldiers who suffered
from bacillary dysentery. None of the latter cats took the complaint and none
showed changes in the large intestine upon sectioning. The injection of various
bacteria, obtained from a stool of amœbic enteritis pathogenic to cats, remained
without result in both the cats employed for this experiment. Lastly, two cats,
which had been kept with those artificially infected, were taken ill spontaneously
and suffered from the disease. In the opinion of Harris, who ascertained the
harmless nature of bacteria derived from the intestinal flora containing dysenteric
amœbæ, young dogs are capable of being infected.
Within the large intestine an active increase of Entamœba histolytica must occur.
Nevertheless, Jürgens did not definitely find changes that might be interpreted in
this sense. Schaudinn (1903) observed division and gemmation in vivo. Both
processes, in which the nucleus divides by amitosis, can only be distinguished by
the fact that the daughter individuals are similar in binary fission but dissimilar in
gemmation, whether they make their appearance singly or in greater numbers.
Schizogony, resulting in the formation of eight individuals, which is so
characteristic for Entamœba coli, was not observed. (But schizogony, into four
merozoites, is now known to occur. Gemmation processes are apparently
degenerative.)
Resistant stages, which serve for transmission to other hosts, are according to
Schaudinn 17 first formed when the diseased portions commence to heal, or more
accurately, the recovery commences when the vegetative increase of the amœbæ
in the intestine discontinues. The so-called spores of E. histolytica were
distinguished very definitely from those of E. coli; they were said to consist of
spheres of only 3 to 7 µ in diameter, which were surrounded by a double
membrane, at first colourless, but becoming a light brownish yellow colour after a
few hours, and possessing a protoplasmic content containing chromidia. They
were said to arise by fragments of chromatin passing outwards from the nucleus
of the amœba into the surrounding cytoplasm (fig. 9, a) and undergoing so
marked an increase that finally the whole cytoplasm became filled with chromidia.
The remainder of the nucleus underwent degeneration and became extruded. On
the surface of the cytoplasm there then arose small protuberances containing
chromidia. These processes had been observed in the living organisms. They
gradually divided and separated from membranes which later became yellow. The
remainder of the amœba perished. Craig 18 had also seen phases of this process of
development. It must be remarked that, according to recent researches, these
processes of exogenous sporulation are degenerative in character (see p. 41). The
small spores may be fungi. The “sporulation” processes are only mentioned here
as a warning. They are now only of historic interest. By means of an experiment
made on a cat, Schaudinn ascertained that ingestion of permanent cysts, which
resist desiccation, is the cause of the infection. The animal took food containing
dry fæces with amœba cysts; these fæces came from a patient suffering from
amœbic enteritis in China. On the evening of the third day the cat evacuated
blood-stained mucous fæces which contained large numbers of typical Entamœba
histolytica. On the fourth day after the infection the animal experimented upon
died, and the large intestine showed the changes previously stated.
E. histolytica also is found in the large intestine. This was originally shown to be
the case by Kartulis, and the fact has recently been confirmed from many
quarters. It is also present in the metastatic abscesses of which it is the cause (cf.
among other authors, Rogers, Brit. Med. Journ., 1902, ii, No. 2,177, p. 844; and
1903, i, No. 2,214, p. 1315).
It should lastly be pointed out in this connection that mixed infections also take
place. For instance, in addition to E. histolytica, E. coli, and, under certain
circumstances, flagellates may be found together. In the same way E. coli may
come under observation even in bacillary dysentery. On the other hand, Schaudinn
stated that in cases of dysentery endemic in Istria, Entamœba coli, if it had
hitherto been present, disappeared, to return again after recovery from the illness.

Fig. 8.—Entamœba histolytica. a, trophozoite (tetragena type)

containing red blood corpuscles, × 1,300; b and c, two isolated
nuclei showing different appearances of karyosome, centriole and
nuclear membrane, × 2,600. (After Hartmann.)

(Entamœba tetragena, Viereck, 1907.)

This amœba must now be considered to be a part of the lifecycle of
Entamœba histolytica, in fact a very important part of that cycle,
especially in its tetranucleate cystic stages.
This organism, the so-called Entamœba tetragena, may occur in the
human intestine in cases of amœbic dysentery, especially in mild or
chronic cases. It was discovered by Viereck in 1907 in patients
suffering from dysentery contracted in Africa. Soon afterwards an
independent description was published by Hartmann, who called the
amœba E. africana. It was also studied by Bensen and Werner.
Recently (1912–13) much work has been published on this amœba
by Darling and others; in this way its relationship to Schaudinn’s E.
histolytica has been made known.
In general morphology it somewhat resembles Entamœba coli, and
its discoverer at first mistook it for a variety of that species.
According to Hartmann, a distinct ectoplasm is only clearly visible
when a pseudopodium is protruded (fig. 5). The granular endoplasm
may contain ingested red blood corpuscles (fig. 6). The large, round
nucleus is visible in the fresh state (fig. 8, a). So-called chromidial
masses (? crystalloidal substances) may occur in the cytoplasm.

Fig. 9.—Entamœba histolytica (tetragena form). a, emission of

chromatin from nucleus; b, nuclear division; c, degenerating form
with two nuclei; d, e, f, cysts containing one, two and four nuclei
 respectively, and showing chromidial blocks. × 2,000. (After
Hartmann.)

Some investigators, as Hartmann, 19 lay stress on the internal

structure of the nucleus (fig. 8, b, c), best seen in preparations fixed
wet and stained with iron-hæmatoxylin. The nucleus is limited by a
well-marked nuclear membrane, on the inside of which granules or
nodules of chromatin may occur. There is a karyosome, which, in
successfully stained specimens, shows, at times, a central dot called
a centriole. (The nucleus of Entamœba coli does not contain such a
centriole.) However, the structure of the nucleus varies at different
periods during the life-cycle.
The diameter of the trophozoites or vegetative forms (fig. 8, a) is
variously given as from 20 µ to 40 µ. Multiplication proceeds by
binary fission and also by schizogony into four merozoites. 20
Reproduction takes place by endogenous encystment (fig. 9, d-f),
which is preceded by nuclear division into two, reduction and then
autogamy. The interpretation of the latter phenomenon as autogamy
is disputed by some authors. The round cysts, which may measure
12 µ to 15 µ in diameter, contain four nuclei, together with darkly
staining masses of various shapes, the so-called “chromidial blocks”
(fig. 9, f). The cyst-wall of E. histolytica (tetragena) is thinner than
that of E. coli, and the diameter of the cyst is rather less. E.
histolytica has not yet been cultivated.
Infection in man occurs by way of the mouth by the ingestion of
cysts. A patient showing acute symptoms of dysentery is not usually
infective, for he is merely harbouring the large trophozoites, which,
by experiment, have been shown not to be infective to animals
(kittens) when administered by the mouth. The stools of recovered
patients may still contain cysts, and they may thus act as cyst-
carriers or reservoirs of disease by infecting water and soil. The
stools of such cyst-carriers are often solid, and so cysts of E.
histolytica (tetragena) are easily overlooked. Mathis (1913) 21 points
out that healthy carriers of E. histolytica may be found; 8 per cent.
of the natives of Tonkin examined by him were healthy carriers of
cysts.
In return cases, or prolonged untreated cases of entamœbic
dysentery, a generation of smaller trophozoites is associated with, or
replaces the larger ones. In stools they are frequently refractile and
consequently stain slowly intra vitam. These trophozoites are the
“smaller, senile, or pre-cyst generation” of Darling. This pre-cyst
generation is characterized by the presence of blocks of crystalloidal
substance in the cytoplasm, and by the possession of a prominent,
densely stainable karyosome. Darling believes this generation to be
the same as that described by Elmassian as Entamœba minuta. 22
Walker, 23 Darling, 24 Wenyon 25 and others believe that Entamœba
histolytica, which was only seen by Schaudinn in a single case, that
of a Chinaman, is really E. tetragena. Darling states that if the
published illustrations of E. histolytica and of E. tetragena are
collected from the literature and compared, it will be seen that the
writers have been calling E. histolytica the large trophozoites seen in
dysenteric stools. These large trophozoites frequently display no
karyosome, but they can be demonstrated as E. tetragena by animal
inoculation, or by the history of the case. On the other hand, the
illustrations of E. tetragena show that the authors have been dealing
with the small generation or reduced forms (“E. minuta”), which are
the direct descendants of the large trophozoites. If kittens are
inoculated rectally with dysenteric material containing large
trophozoites, the strain may be carried in successive kittens for four
to six transfers. If, on the other hand, kittens are inoculated rectally
with small trophozoites of the pre-cyst generation, the transmission
cannot be carried through more than one or two kittens. Wenyon
has succeeded in maintaining E. tetragena in kittens for several
generations.
In some of the preparations from the last remove, pathological
forms of the trophozoites may be seen. These show abnormal forms
of budding, especially peripherally, such as have been described by
Schaudinn and by Craig as characteristic of E. histolytica.
Schaudinn’s small peripheral, exogenous buds and cysts are thus
explained. Craig has latterly changed his views.
Further, Darling states that tetragena cysts fed by the mouth to
kittens produce bowel lesions in which trophozoites having the
characters of E. tetragena, E. histolytica and E. nipponica (Koidzumi)
occur.
In view of the work of recent observers, the peculiar exogenous
encystment which Schaudinn made characteristic of Entamœba
histolytica has been shown to be due to degenerative changes in
senile races of the amœba. E. histolytica and E. tetragena are one
and the same species, and its trophozoite is subject to variation.
According to some observers the histolytica type of nucleus—
described by Schaudinn as being poor in chromatin and not easily
seen in the fresh state—occurs frequently in patients with severe
symptoms of dysentery; on the other hand, the tetragena type of
nucleus—round and easily seen in the fresh state—may occur in
cases presenting slight dysenteric symptoms. Intermediate types of
nuclei are seen. The name of this species, the principal pathogenic
amœba of man, must then be E. histolytica by priority. The cystic
stages of E. histolytica are those first recorded by Viereck and
formerly described as E. tetragena. The geographical distribution of
E. histolytica is wide.

Noc’s Entamœba (1909).

A species of Entamœba was cultivated by Noc 26 in 1909 from cysts
derived from liver abscesses, from dysenteric stools and from the
water supply of Saigon, Cochin China. He cultivated it in association
with bacteria. It is pathogenic. It has been considered allied to E.
histolytica, and shows internal segmentation or schizogony. It
exhibits polymorphism. This amœba has been found by Greig and
Wells (1911) in cases of dysentery in India. It is an important
organism and requires further investigation.
Certain other Entamœbæ 27 have been described at various times
from the intestinal tract of man. Probably most, if not all, of these
are not good species and in some cases much more information is
needed.
Entamœba tropicalis (Lesage, 1908). This parasite is said to be non-
pathogenic, and to occur in the intestine of man in the tropics. It has
a general resemblance to E. coli, but forms small cysts (6 µ to 10 µ
in diameter). The nucleus of the cyst is said to break up into a
variable number of daughter nuclei, from three to thirteen having
been noted. Lesage states that it is culturable in symbiosis with
bacteria. It is probably a variety of E. coli, if not a cultural amœba.
Entamœba hominis (Walker, 1908) has a diameter of 6 µ to 15 µ. A
contractile vacuole is present. Encystment is total, and small cysts
are formed. It is culturable. The original strain, now lost, was
obtained from an autopsy in Boston Hospital. This organism is
probably a cultural amœba.
Entamœba phagocytoides (Gauducheau, 1908). This parasite was
discovered in a case of dysentery at Hanoi, Indo-China. The amœba
is small, 2 µ to 15 µ in diameter. It is active. It ingests bacteria and
red blood corpuscles, while peculiar spirilla-like bodies are found in
its cytoplasm. It multiplies by binary and multiple fission. It can be
cultivated. More recently (1912) the author appears to consider the
amœba to be a stage of a Trichomonas, but abandons the view later
(1914). Further researches on this organism are needed.
Entamœba minuta (Elmassian, 1909) 28 was found, in association
with E. coli, in a case of chronic dysentery in Paraguay. It resembles
E. tetragena but is smaller, rarely exceeding 14 µ in diameter.
Schizogony occurs, four merozoites being produced. The encystment
is total and endogenous, giving rise to cysts containing four nuclei.
This amœba is considered by Darling and others to be the pre-cyst
trophozoite stage of E. histolytica (tetragena).
Entamœba nipponica (Koidzumi, 1909) was found in the motions of
Japanese suffering from dysentery or from diarrhœa, in the former
case in company with Entamœba histolytica. Its diameter is 15 µ to
30 µ. The endoplasm is phagocytic for red blood corpuscles. The
nucleus is well defined, resembling that of E. coli and of E.
tetragena. Multiplication occurs by binary fission and by schizogony.
Encystment is total, but has not been completely followed. Darling
and others consider that this is an abnormal form of E. histolytica,
while Akashi (1913) doubts if it is an amœba at all, but rather is to
be regarded as shed epithelial cells.
General Remark.—It is now considered by some workers that true
Entamœbæ cannot be cultivated on artificial media. Quite recently
Williams and Calkins (1913) 29 have somewhat doubted this opinion,
and state that certain cultural amœbæ, originally obtained from
Musgrave in Manila, exhibit the various morphological variations
associated with true entamœbæ of the human digestive tract.

Entamœba buccalis, Prowazek, 1904.

The size varies from 6 µ to 32 µ. Ectoplasm is always present; the
endoplasm contains numerous food-vacuoles. The nucleus is
vesicular, with a greenish tinted membrane which is poor in
chromatin. The size of the nucleus is from 1·5 µ to 4·5 µ. A
contractile vacuole is not visible. The pseudopodium is broad. It was
discovered in the mouths of persons with dental caries at Rovigno
and also at Trieste, being most easily found in dense masses of
leucocytes, also among leptothrix and spirochæte clusters. It can be
easily distinguished from leucocytes by more intense staining with
neutral red. Multiplication proceeds by fission. Transmission may
take place through the small spherical cysts. This species (fig. 10)
has since been observed in Berlin, and is also occasionally found in
carcinoma of various regions of the oral cavity. (Leyden and
Löwenthal, 1905).

Fig. 10.—Entamœba buccalis, Prow. a-d, the same specimen

observed during five minutes. × 1,000. e, amœba fixed and stained
with iron-hæmatoxylin. × 1,500. (After Leyden and Löwenthal.)

Entamœba buccalis, Prow., is said to be allied to a protozoön which

A. Tietze has found either encysted or free in the lumen of the
orifice of the parotid gland of an infant aged 4 months. The gland
had undergone pathological change, and had therefore been
extirpated. The organisms, which were roundish and three to four
times the size of the normal epithelial cells of the gland, were
without a membrane and possessed a nucleus in which the
chromatic substance appeared to be contained in a karyosome. Bass
and John’s 30 (Feb. 1915) and Smith, Middleton and Barrett (1914)
state that E. buccalis is the cause of pyorrhœa alveolaris.

Entamœba undulans, Aldo Castellani, 1905.

Under this name a protozoön is described which A. Castellani found in addition to
Entamœba histolytica and Trichomonas intestinalis in the fæces of an European
planter living in Ceylon, who had suffered from amœbic enteritis and liver abscess.
The shape of the body was roundish or oval, 25 µ to 30 µ in the greatest diameter.
It was without a flagellum, but with an undulating membrane, and capable of
protruding a long pseudopodium from different parts of its body at short intervals.
The nucleus could not always be recognized in life; it was, however, always
demonstrable by staining. One or two contractile vacuoles were present. The
protoplasm was finely granular, showing no differentiation into ecto- and endo-
plasm. According to Braun, in spite of the author declaring himself expressly
against the flagellate nature of the parasite, such a nature may be assumed to be
tolerably certain in view of the description and illustration.
It is now considered that Entamœba undulans is a portion of a flagellate, namely,
Trichomonas.

Entamœba kartulisi, Doflein, 1901.

Doflein gave this name to amœbæ, from
30 µ to 38 µ in diameter, which Kartulis
(1893) found on examining the pus of an
abscess in the right lower jaw of an Arab,
aged 43, and in a portion of bone that had
been extracted. The movements of the
amœbæ (fig. 11) were more active than
those of “dysenteric amœbæ.” Their coarsely Fig. 11.—Entamœba
kar­tu­lisi, Dofl., from the
granular cytoplasm contained blood and pus
pus of an ab­scess in the
corpuscles, and a nucleus was generally only lower jaw, show­ing dif­‐
recognizable after staining. Vacuoles were fer­ent stages of move­‐
not seen with certainty. Flexner reported ment. (After Kartulis.)
upon a similar case, and Kartulis published
five additional cases. As in these cases dental caries was present the
infection is likely to have proceeded from the oral cavity as a result
of the carious teeth. Craig 31 (1911) considers that this parasite is
probably identical with Entamœba histolytica.
In the literature the following species have been reported as
occurring in the oral cavity of man:—

Amœba gingivalis, Gros, 1849. [? identical with Entamœba buccalis.]

Amœba buccalis, Sternberg, 1862.
Amœba dentalis, Grassi, 1879.
Far too little, however, is known concerning these to regard them as definite
species, that is, independent organisms; Grassi thinks it even possible there may
have been a confusion in their case with salivary corpuscles. If they really are
amœbæ they are all of them probably identical with Entamœba buccalis.

Genus Paramœba, Schaudinn, 1896.

Schaudinn established the genus Paramœba for a marine rhizopod which
multiplied by division, became encysted at the end of its vegetative life and then
segmented into swarm bodies with two flagella. These multiplied by longitudinal
fission, and finally passed into the condition of Amœbæ. Whether the human
parasite described by C. F. Craig (1906) as

Paramœba hominis.
belonged to this genus was for a time uncertain. It is now placed in a new genus
Craigia, Calkins, 1912, since it possesses only one flagellum. 32
In the amœbic stage it is 15 µ to 25 µ in diameter; ecto- and endo-plasm during
rest are indistinguishable. The body substance is granular, with a spherical, sharply
contoured nucleus and an accessory nuclear body. No vacuoles are present, but
occasionally the endoplasm contains red blood corpuscles. The pseudopodia are
hyaline, finger- or lobe-shaped, and are protruded either singly or in twos.
Multiplication is by binary fission and by the formation of spherical cysts (15 µ to
20 µ in diameter) in which occurs successive division of the nuclei, ultimately
forming ten to twelve roundish bodies each of which soon develops a flagellum.
The flagellate stages have similarly a spherical shape and attain a diameter of 10 µ
to 15 µ. They also occasionally contain red blood corpuscles and pass either
directly or after longitudinal division into the amœboid phase.
Craig found these Amœbæ and the flagellate stage belonging to them in six
patients in the military hospital at Manila (Philippine Islands), five of whom were
suffering from simple diarrhœa whilst the sixth exhibited an amœbic enteritis and
contained also Paramœba hominis, with Entamœba histolytica, Schaudinn. In one
of the other cases, Trichomonas intestinalis was present.

B. Amœbæ from other Organs.

Entamœba pulmonalis, Artault, 1898.

Artault 33 discovered a few amœboid forms with nucleus and vacuole
in the contents of a lung cavity. In the fresh condition they were
distinguishable from leucocytes by their remarkable capacity of light
refraction. They were also much slower than the latter in staining
with methylene blue or fuchsine. Their movements became more
lively in a strong light. Water and other reagents killed them, and
then, even when stained, they could not be distinguished from
leucocytes. They have also been seen by Brumpt. R. Blanchard
found amœbæ which may belong here in the lungs of sheep. A.
pulmonalis is perhaps the same as Entamœba buccalis. Smith and
Weidman 34 (1910, 1914) described an entamœba, E. mortinatalium,
from the lungs and other organs of infants in America.

Amœba urogenitalis, Baelz, 1883.

This species was found in masses in the sanguineous urine as well
as in the vagina of a patient in Japan, aged 23. Shortly before the
death of the patient, which was caused by pulmonary tuberculosis,
hæmaturia with severe tenesmus of the bladder had set in. The
amœba, which showed great motility, and had a diameter of about
50 µ when quiescent, exhibited a granular cytoplasm and a vesicular
nucleus. Baelz is of opinion that these parasites were introduced into
the vulva with the water used for washing the parts, and thence had
penetrated into the bladder and vagina. Doflein places the organism
in the genus Entamœba, and it is perhaps identical with E.
histolytica.
Similar cases are also reported (1892–3) by other authors: Jürgens, Kartulis,
Posner, and Wijnhoff. Jürgens found small mucous cysts, filled with amœboid
bodies, in the bladder of an old woman suffering from chronic cystitis; they were
also found in the vagina. The amœba observed by Kartulis in the sanguineous
urine of a woman, aged 58, suffering from a tumour of the bladder, measured
12 µ to 20 µ, and exhibited slow movements by protruding short pseudopodia.
The vacuoles and nucleus became visible only after staining with methylene blue.
Posner’s case related to a man, aged 37, who had hitherto been quite healthy and
had never been out of Berlin. Suddenly, after a rigor, he passed urine tinged with
blood. This contained, besides red and white blood corpuscles and hyaline and
granular casts, large granular bodies (about 50 µ in length and 28 µ in breadth),
which slowly altered their shape, and contained red blood corpuscles in addition to
other foreign matter. These bodies exhibited one or several nuclei and some
vacuoles. From the course of the disease, which extended over a year, and during
which similar attacks recurred, Posner came to the conclusion that the amœbæ
which had originally invaded the bladder had penetrated into the pelvis of the
kidney, where they probably had settled in a cyst, and thence induced the
repeated attacks.
Wijnhoff observed four cases of amœburia in Utrecht.

Amœba miurai, Ijima, 1898.

Under this term the author describes protoplasmic
bodies which Miura, in Tokyo, found in the serous
fluid of a woman, aged 26, who had died from
pleuritis and peritonitis endotheliomatosa. Two days
before death these same forms had also appeared in
the hæmorrhagic fæces of the patient. The bodies
were usually spherical or ellipsoidal, and at one pole Fig. 12.—Amœba miurai, Ij.
carried a small protuberance (fig. 12) beset with × 500. a, fresh; b, after
filamentous short “pseudopodia” (really a treat­ment with di­lute acet­ic
pseudopodium covered with cilia). Their size varied acid. (After Ijima.)
between 15 µ and 38 µ. The cytoplasm was finely
granular, and no difference was observable in the ecto- and endo-plasm, only the
villous appendage was clearer. The cytoplasm contained vacuoles more or less
numerous, none of which was contractile. After the addition of acetic acid one to
three nuclei could be distinguished, 8 µ to 15 µ in size. Actual movements were
not observed. Taking everything into consideration, the independent nature of
these bodies is, to say the least, doubtful, although it cannot be denied that they
possess a certain similarity to the marine Amœba fluida, Grüber or Greeff, and to a
few other species. (It is likely that cells present in serous exudation were mistaken
for amœbæ.)

Appendix.
“Rhizopods in Poliomyelitis acuta.”
In three cases of poliomyelitis acuta which were investigated by Ellermann, the
spinal fluid obtained by puncture of the cord contained bodies, from 10 µ to 15 µ
in size, which had amœboid movements and exhibited variously shaped
pseudopodia in large numbers. After staining, a usually excentric nucleus, about
1·5 µ in size, was demonstrated in them.

Order. Foraminifera, d’Orbigny.

The order is divided by Max Schultze into Monothalamia and Polythalamia. Only a
few of the former can be considered here.

Sub-Order. Monothalamia. (Testaceous Amœbæ).

These forms occur frequently in fresh water, rarely in sea water.
They possess a shell which is either pseudo-chitinous in character, or
consists of foreign particles, or in a few cases is composed of
siliceous lamellæ. There is usually an orifice for the protrusion of
pseudopodia. The only representative of the order of interest here
is:—

Genus. Chlamydophrys, Cienkowski, 1876.

The genus is based on a form which A. Schneider carefully investigated and
considered to be the Difflugia enchelys of Ehrenberg. L. Cienkowski rediscovered
this same form and created for it the genus Chlamydophrys. We agree with this
view, but not with the renaming of the organism (so common at the time). If the
parasite in dung, Chlamydophrys stercorea Cienk. is identical with Difflugia
enchelys of Ehrenberg, the old specific name should be retained.

The genus is characterized by the possession of a hyaline,

structureless, slightly flexible shell which is ovoid or reniform. At the
more pointed pole there is an orifice situated terminally or somewhat
laterally, serving for the emergence of the filiform pseudopodia
(fig. 13, a). The protoplasm does not entirely fill the interior of the
shell. An equatorial zone bearing excretory granules divides the shell
internally into two almost equal portions. The anterior portion is rich
in vacuoles and serves for the reception of nutriment and for
digestion. The posterior part is vitreous, and contains the nucleus.
One to three contractile vacuoles are situated in the equatorial zone.

Chlamydophrys enchelys, Ehrbg.

Syn.: Chlamydophrys stercorea, L. Cienkowski.

This species (fig. 13) is found in the fæces of various animals (cattle,
rabbits, mice, and lizards), and also in quite fresh human fæces.
According to Schaudinn, the parasite occurs so frequently in the
human fæces that it must be considered of wide distribution. The
species must traverse the intestine of man and animals during one
stage of its life cycle, as Schaudinn showed by experiments on
himself and on mice. He infected himself with cysts (fig. 14) by
swallowing them, and evacuated the first Chlamydophrys as early as
the following day. After the evacuation of numerous specimens on
one of the following days the infection ceased.
The nucleus of a living specimen is surrounded by a hyaline, strongly
refractile chromidial mass, arranged in the form of a ring. Chromatin
stains colour it darkly.
Asexual multiplication (fig. 13, b), which takes place in fæces,
follows a similar course to that of allied forms (e.g., Euglypha,
Centropyxis). It commences by the cytoplasm issuing from the
orifice of the shell and assuming the shape characteristic of the
mother organism, but in a reverse position. The nucleus then divides
by mitosis, when the daughter nuclei move apart from one another.
The chromidial ring also divides into two portions by a process of
dumb-bell like constriction. The one daughter nucleus remains in the
mother organism, the other moves towards the daughter individual,
which then separates from the parent.
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