Evolution of parasitism in mammal-associated mites of the group Psoroptidia (Acari:Astigmata)

Host-parasite relationships of mammals and astigmatan mites (Acariformes: Astigmata) belonging to the parvorder Psoroptidia are analyzed. The absolute majority of mammal-associated psoroptidians belongs to the paraphyletic superfamily Sarcoptoidea. Mites of the family complex Psoroptidae (Lobalgidae, Psoroptidae, and Paracoroptinae) shifted from birds to placental mammals independently from each other. Mites of the family complex Sarcoptidae, including all other sarcoptoid families, derived from the common stalk of Psoroptidia independently from the Psoroptid complex. Mites of the sarcoptid complex shifted from nidicoly in mammalian nests to the permanent parasitism on these hosts. They are widely represented on both marsupial and placental mammals and are absent on Monotremata.     

Origin and higher-level relationships of psoroptidian mites (Acari: Astigmata: Psoroptidia): evidence from three nuclear genes

Phylogenic relationships of the Psoroptidia, a group of primarily parasitic mites of vertebrates, were investigated based on sequences from three nuclear genes (4.2 kb aligned) sampled from 126 taxa. Several morphological classification schemes and a recent molecular analysis, suggesting that the group may not be monophyletic were statistically rejected by newly generated molecular data, and the results are robust under a range of analytical and partition strategies. Six families Psoroptidae, Lobalgidae (mammalian parasites), Pyroglyphidae (house dust mites and parasites inside feather calamus), Turbinoptidae (upper respiratory track parasites of birds), Psoroptoididae (downy feather mites), and Epidermoptidae (skin parasites of birds) form a well-supported monophyletic group (the epidermoptid-psoroptid complex). These relationships, recovered by combined and separate analyses of all gene partitions, were previously suspected based on some morphological evidence, but evidence has been dismissed as resulting from convergence based on similar parasitic ecologies. The existence of the epidermoptid-psoroptid complex and the statistical rejection of Sarcoptoidea (the morphology-based group joining all mammal-associated mites) indicate that current classification criteria, influenced as they are by host preferences, need to be reassessed for non-pterolichoid superfamilies. However, two of our findings remain sensitive to analytical methods and assumptions: (i) the families Heterocoptidae and Hypoderatidae as the first and second closest outgroups of Psoroptidia, respectively, and (ii) the superfamily Pterolichoidea (including Freyanoidea) forming a sister clade to the remaining psoroptidian superfamilies. Our findings suggest that (i) house dust mites (Pyroglyphidae: Dermatophagoidinae) originated from a parasitic ancestor within the core of Psoroptidia, violating a basic principle of evolution that it is virtually impossible for a permanent parasite to become free-living, and (ii) there were at least two shifts from presumably avian to mammalian hosts.     

Cebus apella (Primata: Cebidae) as a new host for Fonsecalges johnjadini (Acari: Psoroptidae, Cebalginae) with a description of anatomopathological aspects

Mites collected from the auditory canal of Cebus apella (capuchin monkey), family Cebidae, were identified as Fonsecalges johnjadini (Psoroptidae, Cebalginae). It is the first record of this parasite from this monkey. This paper emphasizes the importance of clinical and anatomopathological examinations for parasitic diagnosis in wild animals.     

Morphological adaptations of acariform mites (Acari: Acariformes) to permanent parasitism on mammals

The external morphological adaptations to parasitism in acariform mites (Acari: Acariformes), permanently parasiting mammals, are briefly summated and analyzed. According to several external morphological criteria (structures of gnathosoma, idiosoma, setation, legs and life cycle), the following six morphoecotypes were established: skin mites (i)-- Cheyletidae, Chirorhynchobiidae, Lobalgidae, Myobiidae, Myocoptidae (the most part), Rhyncoptidae, Psoroptidae; fur mites (ii)--Atopomelidae, Clirodiscidae, Listrophoridae, Myocoptidae (Trichoecius only); skin burrowing mites (iii)--Sarcoptidae; intradermal mites (iv) - sorergatidae and Demodicidae; interstitial mites (v) - pimyodicidae; respiratory mites (vi) - reynetidae, Gastronyssidae, Lemurnyssidae, Pneumocoptidae. In the case of prostigmatic mites, the detailed reconstruction of the origin and evolution of "parasitic" morphoecotypes is possible due to the tentative phylogenetic hypotheses, which were proposed for the infraorder Eleutherengon, a, including the most part of the permanent mammalian parasites among prostigmatic mites (Kethley in Norton, 1993; Bochkov, 2002). The parasitism of Speleognathinae (Ereynetidae) in the mammalian respiratory tract arose independently of the other prostigmats. It is quite possible that these mites switched on mammals from birds, because they are more widely represented on these hosts than on mammals. The prostigmatic parasitism on mammalian skin seems to be originated independently in myobiids, in the five cheyletid tribes, Cheyletiellini, Niheliini, and Teinocheylini, Chelonotini, Cheyletini, and, probably, in a cheyletoid ansector of the sister families Psorergatidae-Demodicidae (Bochkov, Fain, 2001; Bochkov, 2002). Demodicids and psorergatids developed adaptations to parasitism in the skin gland ducts and directly in the epithelial level, respectively in the process of the subsequent specialization. Mites of the family Epimyodicidae belong to the phylogenetic line independent of other cheyletoids. These mites possess the separate chelicerae and, therefore, can not be included to the superfamily Cheyletoidea. It is not quite clear whether they were skin parasites initially or they directly switched to parasitism from the predation. The phylogeny of sarcoptoid mites (Psoroptidia: Sarcoptoidea) is not developed, however, some hypotheses about origin and the following evolution of their morphoecotypes can be proposed. We belive that astigmatic mites inhabiting the mammalian respiratory tract transferred to parasitism independently of other sarcoptoids. The idiosoma of these mites is not so much flattened dorso-ventrally and has proportions which are similar to hose of free-living astigmatids. Moreover, in the most archaic species, the legs are not shortened or thickened as in the most parasites. The disappearance of many morphological structures in these mites, probably, happened parallely with some other sarcoptoids due to their parasitic mode of life. The skin inhabiting sarcoptoids belong to the "basic" morphoecotype, and all other sarcoptoid morphoecotypes, excluding respiratory mites, are derived from it. Some mites of this morphoecotype live on the concave surfaces of the widened spine-like hairs of the rodents belonging to the family Echimyidae (mites of the subfamily Echimytricalginae), in the mammalian ears (some Psoroptidae) or partially sink into the hair follicles (Rhynocoptidae). Finally, mites of the family Chirorhynchobiidae live on the bat wing edges attaching to them by their "ixodid-like" gnathosoma. The fur-sarcoptoids, probably, originated from the skin mites. This morphoecotype is divided onto two subtypes: mites with the dorso-ventrally flattened idiosoma (subtype I) and mites with the teretial idiosoma (subtype II). Each "fur-mite" family includes mites of the both subtypes. All mites of the first subtype belong to the early derivative lineages in their families. Among listrophorids such early derivative lineage is represented by the subfamily Aplodontochirinae (Bochkov, OConnor, 2006), and among Chirodiscidae--by mites of the subfamilies Chirodiscinae, Schizocoptinae, and Lemuroeciinae. Among the "fur" astigmatid families, the family Atopomelidae. probably, is the most archaic, and the most part of atopomelids belongs to the first subtype. However there are several more specialized atopomelid genera belonging to the second subtype, Atopomelus, Dasyurochirus, Lemuroptes, Murichirus, Metachiroecius etc. We believe that mites of the first subtype are represented by the "intermediate" forms between skin mites and mites of the second subtype. Some skin sarcoptoids transferred from skin parasitism to burrowing of the host skin (Sarcoptidae). The established morphoecotypes are partially corresponding to some morphoecotypes established by Mironov (1987) for feather mites. Our morphoecotypes of skin and skin burrowing mites perfectly correspond to Mironov's epidermoptoid and knemidocoptoid morphoecotypes, respectively. The proctophylloid morphoecotype (mites living on the wing feathers), which is the most widely represented within feather mites, has an analogy among mammalian mites - the subfamily Echimytricalginae. The analgoid (mites living in the down feathers) and dermoglyphoid (mites living in the feather quills) morphoecotypes have no analogues among mammalian mites for the obvious reasons. It is interesting why some microhabitats on the host body are not still occupied by prostigmatic or astigmatic mites. We believe that the nutrition is the main limitative factor here. The parasitic prostigmates evolved from predators and, therefore, feed on content of the living cells. They need the direct contact with the live tissues of the host and they belong, therefore, to the morphoecotypes represented by the respiratory, skin, gland duct, intradermal, and interstitial mites. Whereas, the most part of the skin inhabiting astigmats feed on the dead epithelial scales. For this reason these mites, so easily colonized fur of their hosts and feed on the hair grease there. On the other hand, some sarcoptoids transferred to the true parasitism and feed on the cambial cells of the skin epithelium. More over we do not know exactly about nutrition of rhyncoptids yet.     

The major histocompatibility complex in monotremes: an analysis of the evolution of <Emphasis Type="Italic">Mhc</Emphasis> class I genes across all three mammalian subclasses

We report the isolation and characterization of cDNA clones of expressed, functional major histocompatibility complex class-I ( Mhc-I) genes from two species of monotremes: the duck-billed platypus and the short-beaked echidna. The cDNA clones were isolated from libraries constructed from spleen RNA, clearly establishing their expression in at least this one peripheral lymphoid organ. From the presence of conserved amino acid residues, it appears the expressed sequences encode molecules that likely function as classical Mhc-I. These clones were isolated using monotreme Mhc-I processed pseudogenes as probes. These processed pseudogenes were isolated from genomic DNA and, based on their structure, are likely independently derived in the platypus and echidna. When all the monotreme sequences were included in phylogenetic analyses, we found no apparent orthologous relationships between the platypus and echidna Mhc-I. Analyses that included a large number of Mhc-I sequences from other taxa support a separate monotreme Mhc-I clade, basal to a therian Mhc-I clade that is comprised of sequences from marsupial and placental mammals. The phylogenies also support the hypothesis that Mhc-I genes of placental mammals, marsupials, and monotremes are derived from three separate lineages of Mhc-I genes, best explained by two rounds of duplications and deletions. The first round would have occurred prior to the divergence of monotremes and therians, and the second prior to the divergence of marsupials and placental mammals. The sequences described here represent the first reported functional monotreme Mhc-I, as well as the first processed pseudogenes of any type from monotremes.     

Acaricidal activity of aqueous extracts of camomile flowers, Matricaria chamomilla, against the mite Psoroptes cuniculi

Arcaricidal properties of decoctions, infusions and macerates of dried flower heads of camomile, Matricaria chamomilla L. (Asterales: Asteraceae) were tested in vitro against the mite Psoroptes cuniculi Delafond (Parasitiformes: Psoroptidae). This mite species is responsible for otoacariasis in domestic animals. Mites were exposed to the extracts for 24, 48 or 72 h. All the extracts tested showed highly significant acaricidal activity when compared with controls. Among them, a decoction of 10% was the only formulation which gave 100% activity at all the three observations times.     

Origins and divergence times of mammalian class II MHC gene clusters

The class I and II major histocompatibility complex (MHC) genes are apparently subject to evolution by a birth-and-death process. The rate of gene turnover is much slower in the latter genes than in the former. In placental mammals, the class II region can be subdivided into different orthologous subregions or gene clusters (DR, DQ, DO, and DN), but the origins and evolutionary relationships of these gene clusters are not well established. Here we report the results of our study of the times of origin and evolutionary relationships of these gene clusters in mammals. Our analysis suggests that both class II alpha-chain and beta-chain gene clusters are shared by placental mammals and marsupials, but the gene clusters from nonmammalian species are paralogous to mammalian gene clusters. We estimated the times of divergence between gene clusters in placental mammals using the linearized tree and distance regression methods. Our results indicate that most gene clusters originated 170-200 million years (MY) ago, but that DO beta-chain genes diverged from the other beta-chain gene clusters approximately 210-260 MY ago. The phylogenetic trees for the alpha- and beta-chain genes were not congruent, suggesting that the evolutionary history of the class II gene clusters is more complex than previously thought.     

Insights into the evolution of mammalian telomerase: Platypus TERT shares similarities with genes of birds and other reptiles and localizes on sex chromosomes

ABSTRACT: BACKGROUND: The TERT gene encodes the catalytic subunit of the telomerase complex and is responsible for maintaining telomere length. Vertebrate telomerase has been studied in placental mammals, fish, and the chicken, but less attention has been paid to other vertebrates. The platypus occupies an important evolutionary position, providing unique insight into the evolution of mammalian genes. We report the cloning of a platypus TERT (pTERT) ortholog, and provide a comparison with genes of other vertebrates. RESULTS: The pTERT encodes a protein with the high homology to marsupial TERT and avian TERT. Like the TERT of sauropsids and marsupials, as well as that of sharks and echinoderms, pTERT contains extended variable linkers in the N-terminal region suggesting that they were present already in basal vertebrates and lost independently in placental mammals and ray-finned fish. Several alternatively spliced pTERT variants structurally similar to avian TERT variants were identified. Telomerase activity is expressed in all platypus tissues similarly to cold-blooded animals and murine rodents. pTERT was localized on pseudoautosomal regions of sex chromosomes X3/Y2, expanding the homology between human chromosome 5 and platypus sex chromosomes. The synteny analysis suggests that TERT co-localized with sex-linked genes in the last common mammalian ancestor. Interestingly, female platypuses express higher levels of telomerase in heart and liver tissues than do males. CONCLUSIONS: pTERT shares many features with TERT of the reptilian outgroup, suggesting that pTERT represents the ancestral mammalian TERT. Features specific to TERT of eutherian mammals have, therefore, evolved more recently after the divergence of monotremes.     

Structure and function of the gnathosoma of the mange mite,Psoroptes ovis

Mites of the genus Psoroptes (Acari: Psoroptidae) are obligate, non-burrowing, astigmatid ectoparasites of mammals. A detailed understanding of the morphology of the gnathosoma is an important step towards elucidation of the feeding behaviour of this mite and, hence, the pathology of psoroptic mange. Scanning and transmission electron microscopy were used to examine Psoroptes ovis (Hering) (syn. P. cuniculi). The gnathosoma is composed of an infracapitulum (hypostome), which forms the floor of an extended U-shaped pre-oral trough, bounded by laterally compressed palps and enclosing paired chelicerae. Distally, each palp terminates in a flap-like process and four tapered terminal sensillae. The floor of the infracapitulum is interrupted along its length by a dorsally projecting ridge. At the distal end of the infracapitulum are paired, grooved pseudorutellae and, between them, paired, ungrooved lateral lips. Between the lateral lips is a grooved, elongate labium. Medially, a pair of finger-like projections emanates from the paraxial walls of the palps and project anteriorly. Each chelicera is made up of fixed and mobile digits, with a cheliceral membranous fold articulating the two. The distal ends of both digits are chelated. For much of its length, the mobile cheliceral digit appears to lie within a fold formed by the infracapitular ridge, thereby creating a central channel between the chelicerae. It is suggested that this arrangement of elements may facilitate a two-way flow of liquid, where saliva flows down a central salivary canal and spills out onto the host's skin over the labium at the point where the chelated tips of the chelicerae meet and abrade the skin. Liquid food may move along the grooves of the pseudorutellae and then be sucked up a food canal formed by the paraxial walls of the palps and the outer wall of the chelicerae.     

Retroposed elements as archives for the evolutionary history of placental mammals

Reconstruction of the placental mammalian (eutherian) evolutionary tree has undergone diverse revisions, and numerous aspects remain hotly debated. Initial hierarchical divisions based on morphology contained many misgroupings due to features that evolved independently by similar selection processes. Molecular analyses corrected many of these misgroupings and the superordinal hierarchy of placental mammals was recently assembled into four clades. However, long or rapid evolutionary periods, as well as directional mutation pressure, can produce molecular homoplasies, similar characteristics lacking common ancestors. Retroposed elements, by contrast, integrate randomly into genomes with negligible probabilities of the same element integrating independently into orthologous positions in different species. Thus, presence/absence analyses of these elements are a superior strategy for molecular systematics. By computationally scanning more than 160,000 chromosomal loci and judiciously selecting from only phylogenetically informative retroposons for experimental high-throughput PCR applications, we recovered 28 clear, independent monophyly markers that conclusively verify the earliest divergences in placental mammalian evolution. Using tests that take into account ancestral polymorphisms, multiple long interspersed elements and long terminal repeat element insertions provide highly significant evidence for the monophyletic clades Boreotheria (synonymous with Boreoeutheria), Supraprimates (synonymous with Euarchontoglires), and Laurasiatheria. More importantly, two retropositions provide new support for a prior scenario of early mammalian evolution that places the basal placental divergence between Xenarthra and Epitheria, the latter comprising all remaining placentals. Due to its virtually homoplasy-free nature, the analysis of retroposon presence/absence patterns avoids the pitfalls of other molecular methodologies and provides a rapid, unequivocal means for revealing the evolutionary history of organisms.     

Placental endogenous retrovirus (ERV): structural,functional, and evolutionary significance

That endogenous retrovirus (ERV) is present within the placenta of humans and other mammals has been known for the past 25 years, but the significance of this observation is still not fully understood. Much molecular biological data have emerged in recent years to support the earlier electron microscopic data on the presence of placental ERV. The evidence for ERV in animal and human placental tissue is presented, then integrated with data on the the presence of ERV in a range of other tissues, in particular teratocarcinoma cells. Placental invasiveness and maternal immunosuppression are then discussed in relation to metalloproteinase secretion, the immunosuppressive potential of retroviruses, and placental growth factors, while the evidence for a functional link between placental proto-oncogenes and trophoblast malignancy is reviewed. Finally, placental development, structure, and life span are discussed within an evolutionary context. The hypothesis that one or more ancient trophoblastic ERVs could have played a role in the evolution and divergence of all placental mammals is evaluated. BioEssays 20: 307-316, 1998. © 1998 John Wiley & Sons, Inc.     

Genome sequence comparison reveals independent inactivation of the caspase-15 gene in different evolutionary lineages of mammals

We have recently demonstrated that placental mammalian species such as pig and dog express a novel proapoptotic protease, caspase-15, whereas mouse and humans lack this enzyme. Here we investigated the evolutionary fate of the caspase-15 gene in different mammalian lineages by analyzing whole-genome shotgun sequences of 30 mammalian species for the presence of caspase-15 orthologs. Caspase-15 gene sequences were found in representatives of all major mammalian clades except for the superorders Afrotheria (tenrec, rock hyrax, and elephant) and Euarchontoglires (rodents, rabbit, tree shrew, and primates), which either lacked any caspase-15-like sequences or contained mutated remnants of the caspase-15 gene. Polymerase chain reaction screenings confirmed the results of the database searches and showed that the caspase-15 gene is expressed not only in various placental mammals but also in the marsupial, Monodelphis domestica. The observed species distribution implies that caspase-15 has originated in an early ancestor of modern mammals and has been conserved, over more than 180 Myr, in marsupials and many placental mammals, whereas it was independently lost in 2 phylogenetically distant clades of placental mammals, that is, Afrotheria and Euarchontoglires. Our data suggest that the inactivation of the caspase-15 gene was not counteracted by, and may even have been driven by, evolutionary constraints in these clades, and therefore, caution against the uncritical use of gene absence for the inference of phylogenetic relationships.     

Ubiquitous mammalian-wide interspersed repeats (MIRs) are molecular fossils from the mesozoic era.

Short interspersed elements (SINEs) are ubiquitous in mammalian genomes. Remarkable variety of these repeats among placental orders indicates that most of them amplified in each lineage independently, following mammalian radiation. Here, we present an ancient family of repeats, whose sequence divergence and common occurrence among placental mammals, marsupials and monotremes indicate their amplification during the Mesozoic era. They are called MIRs for abundant Mammalian-wide Interspersed Repeats. With approximately 120,000 copies still detectable in the human genome (0.2-0.3% DNA), MIRs represent a 'fossilized' record of a major genetic event preceding the radiation of placental orders.     

The biochemical basis for the conjugation of bile acids with either glycine or taurine

All animals, except for the placental mammals, conjugate their bile acids exclusively with taurine. However, in certain of the placental mammals, glycine conjugates are also found. The basis for the appearance of glycine conjugation among the placental mammals was investigated. The reaction of choloyl-CoA with glycine and taurine, as catalysed by the soluble fraction from guinea-pig liver, had a high affinity for taurine and a poor affinity for glycine. The predominant synthesis of glycine conjugates in the guinea pig can be related to the fact that guinea-pig liver contains an unusually low concentration of taurine and a high concentration of glycine. Rabbits make exclusively glycine conjugates and their livers also contain low concentrations of taurine. However, the biochemical basis for their glycine conjugation is more straightforward than in the guinea pig in that the soluble fraction from rabbit liver has a high affinity for glycine and a poor affinity for taurine. Alternative-substrate-inhibition studies with glycine and taurine in soluble fractions from guinea-pig and rabbit liver revealed that glycine and taurine were mutually inhibitory. This suggests that there is only one enzyme for glycine and taurine conjugation in these tissues. The soluble fractions from bovine liver and human liver also made both glycine and taurine conjugates and evidence is presented that suggests that there is only one enzyme in these tissues too. Even the rat, which excretes mostly taurine conjugates, could make both glycine and taurine conjugates in vitro. However, in contrast with all of the placental mammals studied, the supernatant fraction from liver of the chicken, and other non-mammals, could not make glycine conjugates even in the presence of very high concentrations of glycine.     

Giant Panda Genomic Data Provide Insight into the Birth-and-Death Process of Mammalian Major Histocompatibility Complex Class II Genes

To gain an understanding of the genomic structure and evolutionary history of the giant panda major histocompatibility complex (MHC) genes, we determined a 636,503-bp nucleotide sequence spanning the MHC class II region. Analysis revealed that the MHC class II region from this rare species contained 26 loci (17 predicted to be expressed), of which 10 are classical class II genes (1 DRA, 2 DRB, 2 DQA, 3 DQB, 1 DYB, 1 DPA, and 2 DPB) and 4 are non-classical class II genes (1 DOA, 1 DOB, 1 DMA, and 1 DMB). The presence of DYB, a gene specific to ruminants, prompted a comparison of the giant panda class II sequence with those of humans, cats, dogs, cattle, pigs, and mice. The results indicated that birth and death events within the DQ and DRB-DY regions led to major lineage differences, with absence of these regions in the cat and in humans and mice respectively. The phylogenetic trees constructed using all expressed alpha and beta genes from marsupials and placental mammals showed that: (1) because marsupials carry loci corresponding to DR, DP, DO and DM genes, those subregions most likely developed before the divergence of marsupials and placental mammals, approximately 150 million years ago (MYA); (2) conversely, the DQ and DY regions must have evolved later, but before the radiation of placental mammals (100 MYA). As a result, the typical genomic structure of MHC class II genes for the giant panda is similar to that of the other placental mammals and corresponds to BTNL2∼DR1∼DQ∼DR2∼DY∼DO_box∼DP∼COL11A2. Over the past 100 million years, there has been birth and death of mammalian DR, DQ, DY, and DP genes, an evolutionary process that has brought about the current species-specific genomic structure of the MHC class II region. Furthermore, facing certain similar pathogens, mammals have adopted intra-subregion (DR and DQ) and inter-subregion (between DQ and DP) convergent evolutionary strategies for their alpha and beta genes, respectively.     

Xist and the order of silencing

X inactivation is the mechanism by which mammals adjust the genetic imbalance that arises from the different numbers of gene-rich X-chromosomes between the sexes. The dosage difference between XX females and XY males is functionally equalized by silencing one of the two X chromosomes in females. This dosage-compensation mechanism seems to have arisen concurrently with early mammalian evolution and is based on the long functional Xist RNA, which is unique to placental mammals. It is likely that previously existing mechanisms for other cellular functions have been recruited and adapted for the evolution of X inactivation. Here, we critically review our understanding of dosage compensation in placental mammals and place these findings in the context of other cellular processes that intersect with mammalian dosage compensation.     

A web-database of mammalian morphology and a reanalysis of placental phylogeny

Background  

Recent publications concerning the interordinal phylogeny of placental mammals have converged on a common signal, consisting of four major radiations with some ambiguity regarding the placental root. The DNA data with which these relationships have been reconstructed are easily accessible from public databases; access to morphological characters is much more difficult. Here, I present a graphical web-database of morphological characters focusing on placental mammals, in tandem with a combined-data phylogenetic analysis of placental mammal phylogeny.     

Alpha ganglion cells in mammalian retinae

Retinae from species of six orders of mammals (table 1) were processed by an on-the-slide neurofibrillar staining method to establish whether alpha-type ganglion cells are generally present in placental mammals. Alpha cells of the domestic cat, where they were first defined as a type, are used as a standard of reference. Alpha cells were found in all the twenty species examined; characteristically they have the largest somata and large dendritic fields with a typical branching pattern. In keeping with the common morphology there are inner and outer stratifying subpopulations and therefore a presumptive 'on-centre' and 'off-centre' responsiveness to light. Depending on the species, alpha cells form between 1 and 4% of the ganglion-cell population and their dendritic fields cover the retina three to four times. The morphology of alpha ganglion cells, and many of their quantitative features, are conserved in mammals coming from different habitats and having a wide variety of behaviours. Because it is known different habitats and having a wide variety of behaviours. Because it is known from the cat that alpha ganglion cells have brisk-transient or Y receptive fields it is possible that all placental mammals possess this physiological system.