Terminal associations occur commonly between meiotic homologues of the two smallest (S10, S11) chromosomes in the northern race of Cryptobothrus chrysophorus when they are either heterozygous or homozygous for distal supernumerary heterochromatic segments. A detailed examination of the origin and behaviour of these associations provides convincing evidence that they are non-chiasmate in character and so cannot be explained by either pseudoterminalisation or terminalisation. The same is true of the terminal associations involved in the persistent pseudomultiples that develop between non-homologues of Heteropternis obscurella when one or both of these carry distal heterochromatic segments. In both situations the C-bands involved in such terminal associations are entire and are never interrupted by non-banded material. In Cryptobothrus, similar associations can also develop between centromere regions when these are heterozygous or homozygous for proximal supernumerary heterochromatic segments.
Comparative fluorescence studies on the chromosome of ten species of acridid grasshoppers, with varying amounts and locations of C-band positive heterochromatin, indicate that the only regions to fluoresce differentially are those that C-band. Within a given species there is a marked tendency for groups of chromosomes to accumulate heterochromatin with similar fluorescence behaviour at similar sites. This applies to all three major categories of heterochromatin - centric, interstitial and telomeric. Different sites within the same complement, however, tend to have different fluorescence properties. In particular, centric C-bands within a given species are regularly distinguishable in their behaviour from telomeric C-bands. Different species on the other hand, may show distinct forms of differential fluorescence at equilocal sites. These varying patterns of heterochromatin heterogeneity, both within and between species, indicate that whatever determines the differential response to fluorochromes has tended to operate both on an equilocal basis and in a concerted fashion. This is reinforced by the fact that structural rearrangements that lead to the relocation of centric C-bands, either within or between species, may also be accompanied by a change in fluorescence behaviour.
Atractomorpha similis (2 n=19 ♂, 20 ♀) is a hygrophilous, tropical to temperate, species of pyrgomorphine grasshopper. We have sampled 70 populations covering the known distributional range of this species within Australia. All of them proved to be polymorphic for heterochromatin content as revealed by C-band analysis of embryonic neuroblasts. This polymorphism affects all ten members of the basic haploid set and includes variants involving differences in either the presence or the amount of procentric, interstitial and terminal C-blocks, as well as variation in the occurrence and nature of short arms on otherwise telocentric chromosomes. A majority of these variants appear to result from heterochromatin addition since the presumptive sibling, Atractomorpha australis, like other species of the genus that have been C-banded, is generally depauperate in heterochromatin. The net result of this extraordinary polymorphism is that each chromosome of A. similis exists in 10–50 distinct morphs. Consequently, there is a high level of chromosomal heterozygosity in all populations in terms of the number of heterozygous pairs present within a complement and an even higher level in terms of the total range of karyomorph patterns. There is also a wide range of total heterochromatin content, as measured by the percent of the total chromosome area occupied by C-band material, with values ranging from 13% to 44%. Specific marker chromosomes which predominate in particular geographical areas serve to distinguish six major cytotypes within A. similis. The two most southerly of these cytotypes show a narrower range of heterochromatin content but with higher values which reflect the more general occurrence of substantial terminal C-blocks within them. Finally, the populations from Fraser Island constitute a particularly distinctive cytotype characterised by the least number of morphs, the lowest level of chromosomal heterozygosity and a restricted range of heterochromatin content confined to the lower end of the known distributional spectrum.
The endemic Australian grasshopper Heteropternis obscurella shows considerable variation in respect of both chromosome structure and chromosome behaviour. The structural differences depend upon different patterns of heterochromatin distribution as revealed by C-banding. These involve differences between populations in respect of polytypic variation in the size of paracentromeric C-bands and differences within populations in respect of polymorphisms both for terminal blocks of heterochromatin in autosomes 3 to 8 and a large proximal block of heterochromatin in autosome 7. The behavioural differences stem in part from genotypically determined variation in the chiasma distribution pattern which is markedly localised in a majority of populations but more randomly distributed in populations from the south of Western Australia. Behavioural differences also arise as secondary consequences of the presence of those heterochromatic blocks which occur as polymorphisms. The distal blocks on autosomes 5, 6, 7 and 8 lead to a redistribution of chiasmata to more proximal sites while the proximal block on 7 leads to the virtual abolition of chiasma formation in that bivalent and its replacement by a non-chiasmate mechanism of segregation. This depends upon a persistent proximal heterochromatic association between the pairing partners. The presence of distal C-blocks on bivalents 3 to 8 gives rise to persistent pseudomultiples, formed as a result of heterochromatic associations between these blocks. Such pseudomultiples involve any two or three of these six bivalents, provided they carry distal blocks, and their frequency rises dramatically in the presence of the proximal heterochromatic block on chromosome 7.
C-band patterns have been analysed in embryonic neuroblast chromosomes of 23 Australian species of acridoids. All of them showed paracentromeric C-bands but these varied considerably in size both within and between species. Many of them also showed interstitial C-bands in from 1–5 members of the haploid complement and in two cases (Atractomorpha similis and Genus nov. 95 ochracea) larger numbers of interstitial bands were present. Terminal C-bands were the least common though again when present they could be found in 1–6 members of the complement except in the cases of A. similis and Genus nov. 95 ochracea where still larger numbers occur. In 5 of the 23 species the megameric chromosome pair was distinctively C-banded. The B-chromosomes found in 3 of the species were also strikingly different in C-band characteristics compared to the standard A-chromosomes. Differences in the number of very small chromosomes present in different species clearly cannot be explained in terms of differences in their C-band content. Neither are differences in genome size simply related to differences in the total amount of C-band material indicating that changes in the size of the genome in this group have involved alterations in both eu and heterochromatin content. Finally similar amounts of C-band material may be distributed throughout the complement in very different ways in different species.
Synthetic F1 hybrids between individuals taken from selected populations of the northern (Glenn Innés, Bolivia Hill) and southern (Forbes Creek, Mount Aggie) chromosome races of the grasshopper Cryptobothrus chrysophorus confirm that these two ‘races’ are indeed distinguished by fixed differences in heterochromatin content. These differences affect five of the six medium sized members of the complement (M 4, 5, 6, 8 and 9). Added to this there is a marked change in the character of the remaining medium pair (M 7) which functions as the megameric in this species. — Meiosis in F1 males is characterised by the presence of univalency in from 22–32% of the meiocytes. This appears to be genotypic in causation and may involve from one to three chromosome pairs in any one cell. Laggards produced from such univalents lead to a failure of anaphase separation at either first or second division or at both of them. Consequently from 17–36% of the sperm produced are giant (diploid or tetraploid) in size. Added to this the medium members commonly form end to end associations involving the heterochromatic (northern race) and euchromatic (southern race) terminii. This is a modified form of the heterochromatic end associations which occur between homologous medium pairs in northern populations. The two small pairs (S10 and 11) show comparable behaviour in both races when they carry terminal heterochromatic supernumerary segments. Such associations persist despite the fact that they are non-chiasmate in character. Moreover, despite the asynapsis found in F1 males they produce F2s when allowed to intersib mate, although many of the embryos fail to develop, presumably as a result of genotypic imbalance, or else are aneuploid in constitution. In the F2s that do survive to maturity the chromosome differences which distinguish the northern and southern forms are recombined to give a wide variety of karyotypes in which the observed pairing is much improved and from which F3s were subsequently obtained.
A study of the male meiotic system in two populations of the gekko Phyllodactylus marmoratus (Gray) has shown that both total and interstitial chiasma frequencies vary cyclically throughout the year. This variation is consistent in each population and was observed over a number of years. The total chiasma frequency (an index of the number of terminal chiasmata) has a different form of cyclic variation than does the interstitial chiasma frequency, and it is argued that they are under independent genetic controls. Reproductive studies suggest that only the sperm with the lowest total chiasma frequencies and greatest range of interstitial frequencies are used for fertilization. An experimental approach has shown that prolonged exposure to low temperature produces a significant increase in total chiasma frequency. It is believed that this environmental cue is responsible for the cyclic nature of total chiasma frequency.
The gekko Phyllodactylus marmoratus has at least three distinct chromosome races; 2n=36, 2n=36 ZZ/ZW and 2n=34. Specimens from these races are morphologically distinguishable, have a degree of habitat specialization and occur in a defined distribution. The 2n=36 race found in Eastern Australia is the presumed primordial type. The 2n=34 race occurs in Western Australia and is regarded as a fusion derivative. The 2n=36 ZZ/ZW race, which is only found on the Murray River system in Eastern Australia has a heteromorphic sex chromosome system present in the female. Giemsa banding suggests that this heteromorphism is the result of a pericentric inversion.
Comparative electrophoretic phenotypes of 18 of the 32 species of the lizard genus Varanus have been determined for four proteins. The animals studied were representative of species from Africa, Israel, Southeast Asia and Australia. Malate dehydrogenase (A2) exhibited a single phenotype throughout. Lactate dehydrogenase (B4) showed four distinctive electrophoretic forms which grouped the various subgenera as follows: (1) Polydaedalus, Empagusia (African); (2) Psammosaurus (Israel); (3) three species of Varanus, V. gouldii, V. spenceri, V. mertensi (Australian); (4) Dendrovaranus, Indovaranus (Southeast Asian), other Varanus species, Odatria (Australian). Electrophoretic and previously reported karyotypic data were used to interpret the phylogenetic relationships as well as the mode and direction of evolution of these animals. In particular, the results questioned the reality of the subgenus Varanus as a taxonomic unit, since four distinct karyotypic forms and two LDH-B4 phenotypes were observed for these animals, of which one belongs to another subgenus. Serum albumin and carbonic anhydrase phenotypes were of little use in deciding phenotypic groupings.