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PROGRESS IN PROTOZOOLOGY PROGRESS IN PROTOZOOLOGY
22 B. M. HONIGBERG lar rings, micronemes, and a conoid; also, centrioles were seen associated with dividing nuclei. Sexual "behavior" is an important characteristic in ascertaining the taxonomic position of an organism in the subclass COCCIDIA. Therefore, the elucidation of a presumed sexual phase in the life cycle(s) of piroplasms is of major importance. Me hi horn et al. (1980) indicated that the piroplasms have many characteristics in common with the members of HAEMOSPORINA and that the two assemblages should be considered as "sister groups." According to M a 11 m a n (at the present Congress), however his fine-structural observations of a species of Babesia cultivated in vitro revealed apparent syngamy of gametes, a process unlike that typical of HAEMOSPORINA, but resembling fertilization described from several ADELINA. Furthermore, according to Prof. D e s s e r, the kinetes of the species belonging to the genera Babesia and Theileria are as similar to the sporokinetes of Karyolysus spp. as to the ookinetes of HAMEOSPORINA. The picture is further confused by the study of Theileria annulata (S c h e i n et al. 1975) in which the sexual reproduction is said to involve anisogamy, with filariform microgametes. Some workers still doubt the occurrence of sexual reproduction among the piroplasms. In light of the available data, Prof. Desser concluded that a definite commitment on the taxonomic position of the piroplasms among the APICOMPLEXA should be postponed until the problems outlined in his presentation have been resolved. Remarks of the Discussants Dr. Tamara Beyer and Profs. G a r n h a m and K r y 1 o v made remarks about APICOMPLEXA. The following statement was made by Prof. GARNHAM. The two classification schemes published by the Society of Protozoologists (H on i g b e r g et al. 1964; L e v i n e et al. 1980) were concerned only with the higher taxa, down to suborders, but the interrelationships can be more precisely visualized in the taxa below suborders — the former represents concepts, the latter actual organisms. In view of this, he proposed that a special Round Table Discussion be arranged at some future time, for example at the next Congress. The "phylum" APICOMPLEXA, divided into three "sub-classes," seemed to him natural and fairly neatly to fit into the old term SPO- ROZOA which we reluctantly abandoned; but electron microscopy has revolutionized the territory and has necessitated a new look. APICOM- PLEXA are a unique group in: (a) being obligatorily parasitic and (b) possessing ultramicroscopic structures at the anterior end and a micro- http://rcin.org.pl
PHYLOGENETIC RELATIONSHIPS AMONG PROTOZOA 197 pore near the center. These and other structures regress at different stages of development and in the degree of juvenility in phylogeny. They are best seen in sporozoites, the product of the sexual phase, and function particularly in the invasion of a host cell, which is an important feature of parasitism. Three factors are of importance in the phylogeny of APICOMPLE- XA: 1. The Stage: characteristic organelles may be present at one stage but absent in another (e.g., tubular mitochondria, cf. Vivier); 2. The Host: It is essential to consider the phylogeny of both vertebrate and invertebrate hosts in heteroxenous examples, as this does not necessarily run parallel with the phylogeny of the parasite — unfortunately there are few indications as to the date when the parasite became established; 3. Zoogeography. The presence or absence of certain parasites in various vertebrate groups in different regions of the world may be linked with major geological events such as continental drift in more remote ages and glaciation in more recent. Factors 2 and 3 are not directly related to classification, but undoubtedly shed light on phylogeny. The intervention of a biting arthropod has long been suggested as the path taken in the phylogeny of haemosporidia from coccidia and of certain haemogregarines from adeleids. According to Prof. G a r n h a m, the question of free-living ancestors of APICOMPLEXA has assumed new importance in light of the recent observations on Acanthamoeba and Naegleria, which have illustrated how the host barrier in parasitism can be surmounted. However, ideas about their origin remain highly speculative. We look for three primary characters — a degree of "amoebicity," a flagellate stage, and some indication of sexuality. The members of the suborder Bodonina Hollande, 1952 emend. Vickerman, 1976 have been suggested as the possible ancestors of APICOMPLEXA. However, Prof. Garnham's preference is for RHIZOPODA von Siebold, 1875, some members of which possess at least two of these characters, e.g., Naegleria in the order SCHIZOPYRENIDA. The "monopodial" cylinder found in some members of this group resembles the structure of the motile zygote (ookinete). The best secondary clues are offered by the sexual stages, e.g., the microgamete and the zygote, which certainly are of immense significance in the more recent phylogeny. Prof. G a r n h a m was still uncertain as to how much help will be obtained through the employment of biochemical features, e.g., isoenzymes and nucleic acid series. Dr. BEYER was the second discussant. She found no justification for the doubts expressed by some workers with regard to the validity of the name APICOMPLEXA, and adduced the following arguments in support of her viewpoint: http://rcin.org.pl
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- Page 9 and 10: 183 In his abstract Mignot (1981) s
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- Page 51 and 52: 225 brate cycle in vitro. Further c
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PHYLOGENETIC RELATIONSHIPS AMONG PROTOZOA 197<br />
pore near the center. These and other structures regress at different<br />
stages of development and in the degree of juvenility in phylogeny.<br />
They are best seen in sporozoites, the product of the sexual phase, and<br />
function particularly in the invasion of a host cell, which is an important<br />
feature of parasitism.<br />
Three factors are of importance in the phylogeny of APICOMPLE-<br />
XA: 1. The Stage: characteristic organelles may be present at one<br />
stage but absent in another (e.g., tubular mitochondria, cf. Vivier);<br />
2. The Host: It is essential to consider the phylogeny of both vertebrate<br />
and invertebrate hosts in heteroxenous examples, as this does<br />
not necessarily run parallel with the phylogeny of the parasite — unfortunately<br />
there are few indications as to the date when the parasite<br />
became established; 3. Zoogeography. The presence or absence<br />
of certain parasites in various vertebrate groups in different regions of<br />
the world may be linked with major geological events such as continental<br />
drift in more remote ages and glaciation in more recent. Factors<br />
2 and 3 are not directly related to classification, but undoubtedly<br />
shed light on phylogeny. The intervention of a biting arthropod<br />
has long been suggested as the path taken in the phylogeny of<br />
haemosporidia from coccidia and of certain haemogregarines from<br />
adeleids.<br />
According to Prof. G a r n h a m, the question of free-living ancestors<br />
of APICOMPLEXA has assumed new importance in light of the recent<br />
observations on Acanthamoeba and Naegleria, which have illustrated<br />
how the host barrier in parasitism can be surmounted. However,<br />
ideas about their origin remain highly speculative. We look for three<br />
primary characters — a degree of "amoebicity," a flagellate stage, and<br />
some indication of sexuality. The members of the suborder Bodonina<br />
Hollande, 1952 emend. Vickerman, 1976 have been suggested as the<br />
possible ancestors of APICOMPLEXA. However, Prof. Garnham's<br />
preference is for RHIZOPODA von Siebold, 1875, some members of<br />
which possess at least two of these characters, e.g., Naegleria in the order<br />
SCHIZOPYRENIDA. The "monopodial" cylinder found in some members<br />
of this group resembles the structure of the motile zygote (ookinete).<br />
The best secondary clues are offered by the sexual stages, e.g., the<br />
microgamete and the zygote, which certainly are of immense significance<br />
in the more recent phylogeny. Prof. G a r n h a m was still uncertain<br />
as to how much help will be obtained through the employment<br />
of biochemical features, e.g., isoenzymes and nucleic acid series.<br />
Dr. BEYER was the second discussant. She found no justification<br />
for the doubts expressed by some workers with regard to the validity<br />
of the name APICOMPLEXA, and adduced the following arguments in<br />
support of her viewpoint:<br />
http://rcin.org.pl