CHROMOSOMAL DIFFERENTIATION IN CLARKIA DUDLEYANA

Snow , Richard . (U. California, Davis.) Chromosomal differentiation in Clarkia dudleyana. Amer. Jour. Bot. 47 (4) : 302—309. Illus. 1960.—Clarkia dudleyana (n=9) is a common, colonial annual of the early-summer California flora. Of 275 individuals, derived from 9 natural populations and their garden-grown representatives, 17.1% were heterozygous for reciprocal translocations. Supernumerary chromosomes were also found in about 2% of the plants examined. The translocation heterozygotes are not distributed regularly over the species range but are concentrated near the geographical center of distribution. Most of the populations contained none or only a few heterozygotes, but in one colony 69% of 42 plants sampled were heterozygous. Judging from the meiotic metaphase associations observed, at least 5 different chromosome arrangements are present at this locality. Hybrids between colonies have invariably been translocation heterozygotes, the largest association found in such hybrids being a chain of all 18 chromosomes (a potential ring of 18). No correlation is evident between geographical separation and degree of cytological differentiation. Heterozygotes with smaller rings of 4 or 6 chromosomes, whether from natural populations or resulting from interpopulation hybridization, are highly fertile owing to the regular alternate disjunction of the chromosomes of the rings. In the larger rings of 12 to 18 chromosomes, derived from interpopulation crosses, segregation is much more irregular and leads to high sterility. It is possible that at least in some localities the heterozygotes enjoy a selective advantage over their homozygous sibs. It is also postulated that homozygosity for a particular chromosome arrangement may be selectively favored in a certain habitat, as a result of a position effect attendant upon placing formerly non-linked genes in the same linkage group through reciprocal translocation. The high degree of chromosomal differentiation between some populations of this species suggests that the complex heterozygotes of Oenothera have arisen as a result of hybridization of cytologically differentiated races.     

CYTOGENETIC STUDIES IN CLARKIA,SECTION PRIMIGENIA. II. A CYTOLOGICAL SURVEY OF CLARKIA ARCUATA AND CLARKIA LASSENENSIS

Clarkia arcuata and C. lassenensis are the 2 members of the subsection Flexicaules. Although closely related morphologically, they show very different patterns of chromosomal variability in nature. About 25% of the plants grown from wild seed of C. arcuata, a predominantly cross-pollinating species, were heterozygous for 1 or 2 translocations; such heterozygotes were found in 5 of the 9 populations sampled. An analysis of the chromosome pairing in intraspecific crosses indicated that at least 5 different translocations giving a ring of 4 with the “standard” strain, 2 giving a ring of 6, and 2 giving a ring of 8 are present in nature. No arrangement was found with widespread distribution, and it is impossible to say at present what might be the primitive arrangement of this species. One population was found to contain an inversion, a rearrangement which is very rare in Clarkia at the intraspecific level. In C. lassenensis, a predominantly self-pollinating species, only 6% (3 plants) of a sample of 53 were translocation heterozygotes, and these heterozygotes were found in only 2 of 13 populations. Intraspecific crosses indicated that one chromosome arrangement, the “standard,” was present throughout the species range.     

CYTOGENETIC STUDIES IN CLARKIA,SECTION PRIMIGENIA. I. A CYTOLOGICAL SURVEY OF CLARKIA AMOENA

Snow , Richard . (U. California, Davis.) Cytogenetic studies in Clarkia, section Primigenia. I. A cytological survey of Clarkia amoena. Amer. Jour. Bot. 50(4): 337–348. Illus. 1963.—Clarkia amoena (2n = 14) shows great cytological diversity based on reciprocal translocations. About ½ the plants sampled were heterozygotes; in 2 of the subspecies the frequency was as high as ⅔. The metaphase configurations were varied, including rings of 4, 6, 8, 4 + 4, 6 + 4, and more unusual types forming a chain of 3 + a univalent or a chain of 4 + a univalent. The latter have only 13 chromosomes but are not true monosomies. Some plants with supernumerary chromosomes were found, in addition to a plant with only 12 chromosomes which was derived from a “monosomic” type. From crosses of various homozygous lines to a standard strain, it was found that the standard end arrangement of chromosomes occurs widely and commonly throughout the species. A minimum of 7 different end arrangements, giving a ring of 4 with the standard strain, and 1 giving a ring of 6, account for the chromosome pairing found in interstrain crosses. No doubt more would be found with further testing. Some of the translocations may be components of adaptive polymorphic systems, though there is no proof of this yet. No balanced lethal systems similar to those in the closely related genus Oenothera were found.     

CYTOGENETIC STUDIES IN CLARKIA,SECTION PRIMIGENIA. IV. A CYTOLOGICAL SURVEY OF CLARKIA GRACILIS

Clarkia gracilis (2n = 28) is an allotetraploid which combines genomes from two subsections of section Primigenia. Natural populations consist of plants homozygous for a single chromosome arrangement, which may differ from the arrangements in other populations by one or more reciprocal translocations. One arrangement, the “standard,” was widespread. Cytological observations on plants derived from crosses between populations of C. gracilis, and on triploid plants derived from crosses with C. amoena subsp. huntiana, one of the parents of the tetraploid, were used to determine the end arrangements present. Ten arrangements were identified, and it was found that the standard amoena subgenome of C. gracilis is identical in end arrangement to the previously defined standard arrangement of the diploid C. amoena. Hence a race of C. amoena with this arrangement was involved in the hybridization which gave rise to C. gracilis. Evidence was found that other arrangements of C. amoena have probably been introduced into C. gracilis by subsequent introgressive hybridization. Cytological differences, coupled with differences in morphology, ecology, and distribution indicate that C. gracilis should be subdivided into four subspecies instead of the three presently recognized.     

Environmental predictions and distributional limits of chromosomal taxa in the Australian grasshopperCaledia captiva (F.)

Summary Correlations between environmental parameters and the distribution of four chromosomal taxa in the acridine grasshopperCaledia captiva (F.) have been analyzed. A bioclimate prediction system (BIOCLIM) was used as a model to determine their bioclimatic envelopes and to predict their probable geographic distributions, based on climatic similarities with actual distributions. New, and formerly unsuspected, distribution areas are predicted. Parapatric boundaries are defined that actually coincide with known limits. The general distribution of the four taxa is correlated with rainfall seasonality and a temperature gradient. Their distributional boundaries are mostly in accordance with extreme values in rainfall in an east-west direction and temperature in a north-south direction. A nonmetric multidimensional scaling analysis of one selected narrow parapatric boundary confirms the nature of the geographic correlation with several climatic extreme values. The climatic and vegetation analyses suggest that sharp parapatric distributions can be maintained by gradual environmental influences and that coincidence with abrupt ecotones is not necessary. At least one of the parapatric boundaries seems to be relatively recent in origin (2000–6000 years) and on the basis of the location of extreme climatic values as well as chromosomal and genetic markers it is expected to slowly move further south and east of its present location apparently following a climatically established boundary.     

Genomics of natural populations: Evolutionary forces that establish and maintain gene arrangements in Drosophila pseudoobscura

The evolution of complex traits in heterogeneous environments may shape the order of genes within chromosomes. Drosophila pseudoobscura has a rich gene arrangement polymorphism that allows one to test evolutionary genetic hypotheses about how chromosomal inversions are established in populations. D. pseudoobscura has >30 gene arrangements on a single chromosome that were generated through a series of overlapping inversion mutations with >10 inversions with appreciable frequencies and wide geographic distributions. This study analyses the genomic sequences of 54 strains of Drosophila pseudoobscura that carry one of six different chromosomal arrangements to test whether (i) genetic drift, (ii) hitchhiking with an adaptive allele, (iii) direct effects of inversions to create gene disruptions caused by breakpoints, or (iv) indirect effects of inversions in limiting the formation of recombinant gametes are responsible for the establishment of new gene arrangements. We found that the inversion events do not disrupt the structure of protein coding genes at the breakpoints. Population genetic analyses of 2,669 protein coding genes identified 277 outlier loci harbouring elevated frequencies of arrangement‐specific derived alleles. Significant linkage disequilibrium occurs among distant loci interspersed between regions with low levels of association indicating that distant allelic combinations are held together despite shared polymorphism among arrangements. Outlier genes showing evidence of genetic differentiation between arrangements are enriched for sensory perception and detoxification genes. The data presented here support the indirect effect of inversion hypothesis where chromosomal inversions are favoured because they maintain linked associations among multilocus allelic combinations among different arrangements.     

ANALYSIS OF N-ALKANES FROM THREE SPECIES OF CLARKIA

Sixteen n-alkanes were isolated and identified from the herb Clarkia unguiculata, fourteen from C. exilis and seven from C. tembloriensis. These alkanes ranged from C₂₀H₄₂ to C₃₅H₇₂. In all three taxa, the odd numbered alkanes were generally present in greater quantities than even numbered ones, which is similar to the alkane patterns of Monarda that were observed in our laboratories. Clarkia tembloriensis has a characteristically high percentage (95%) of the C₂₀ alkane.     

PATTERNS OF PUFFING ACTIVITY AND CHROMOSOMAL POLYMORPHISM IN DROSOPHILA SUBOBSCURA. IV. EFFECT OF INVERSIONS ON GENE EXPRESSION

We have observed that, contrary to a common assumption, the puffing patterns manifest in the salivary chromosomes of Drosophila subobscura are modified by chromosomal inversions as well as by genic content. An inversion effect is apparent in the E and A chromosomes of five strains coming from four different natural populations. An effect due to the geographical location of the populations is also detected in the J and O chromosomes. The chromosomal and geographic effects are distinguishable but not contradictory. Indeed, a statistical test using the DK² coefficient of distance shows that, for a given chromosomal arrangement, strains of different geographic origin exhibit puffing patterns significantly different; these patterns are, however, more similar to each other than they are to those of strains carrying different chromosomal arrangements of the same chromosome.     

Chromosomal and allelic variation in Drosophila americana: selective maintenance of a chromosomal cline.

Geographically structured genetic variation, as represented by clines and hybrid zones, offers unique opportunities to study adaptation and speciation in natural populations. A hybrid zone has been reported between Drosophila americana americana and Drosophila americana texana, two taxa that are distinguished solely by the arrangement of their X and 4th chromosomes. In this study, samples of D. americana were collected along a latitudinal transect across the inferred hybrid zone, and the frequency of the alternative chromosomal arrangements is reported. These data illustrate that the alternative chromosomal arrangements are distributed along a shallow cline over a broad geographic region, and that the frequency of the arrangements is tightly correlated with latitude. Allelic variants at 13 RFLP loci in three genes on chromosome 4 exhibit no evidence of association with the cline. Presence of a cline for the chromosomal arrangements, as well as a general absence of geographic structure for variation at these genes, is interpreted as evidence that natural selection is responsible for the maintenance of this chromosomal cline. Furthermore, these results demonstrate that taxonomic subdivision of D. americana is unwarranted, because it exists as a cohesive species that is segregating a chromosomal fusion.     

Variable evolutionary response to regional climate change in a polymorphic species

The evolutionary response to regional and global climate change may vary in widespread polymorphic species, so predicting future genetic responses will require careful tracking of genetic variability in local populations. We surveyed chromosomal inversion polymorphisms in 25 populations of Drosophila robusta, many of which have been sampled repeatedly starting in the 1940s, 50s, and 60s up until 2007, across its range in the USA. Frequencies of some northerly, or cold‐adapted, gene arrangements have declined in the face of increasing temperatures, whereas frequencies of several southern, or warm‐adapted, gene arrangements were positively correlated with increasing temperature changes. Over a finer geographic scale, populations from the west‐central part of the species range from the Ozark Plateau, Ouachita mountains, and eastern Oklahoma showed genetic differentiation between south‐central Ozark and western Ozark/Ouachita regions that has persisted in the face of recent shifts in gene arrangement frequencies. Overall, populations of D. robusta exhibited dynamic genetic changes over time, with some populations shifting chromosome frequencies in just 10–15 years. Some temporal genetic shifts were widespread and significantly correlated with temperature increases, but regions of the genome marked by different gene arrangements have responded in different sections of the species range. In some parts of the species range, chromosome frequencies shifted but were not associated with changing temperatures, showed little or no temporal change, or temporal shifts stopped for temperature sensitive gene arrangements near fixation. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 702–718.     

Monitoring chromosomal polymorphism in Drosophila subobscura over 40 years

The inversion chromosomal polymorphism of Drosophila subobscura is considered to be adaptive as a result of its responses at different time scales to temperature changes. This work reports the longest‐term study of chromosomal polymorphism for a single population of D. subobscura with climatic data from the collecting site itself. The chromosomal analysis of D. subobscura samples collected six times over a 40‐year period at the same location and in the same seasonal interval has revealed the continuous presence of 16 common and six moderately rare chromosomal arrangements through the period. This analysis also corroborates the previously detected negative relationship between the frequencies of the standard (cold‐climate) arrangement on each of its five chromosomes and temperature, as well as between a comprehensive measure of cold adaptation (the total autosomal proportion of standard arrangement) and temperature. These and previous results would support that species harboring cold‐ and warm‐adapted polymorphic chromosomal arrangements, like D. subobscura, can rapidly respond to environmental changes.     

天门冬科黄精族细胞学研究进展

在全面收集和整理黄精族染色体数据的基础上,对国内外有关黄精族各类群间的染色体数目和倍性的变化规律进行了总结,并从染色体的多倍化和非整倍化与系统发育关系和地理分布方面探讨了黄精族内各属的起源和演化关系问题。黄精族包括黄精属、舞鹤草属、异黄精属和竹根七属,共约100余种,其中舞鹤草属(x=18)、异黄精属(x=16)和竹根七属(x=20)的染色体基数稳定,而黄精属染色体基数波动较大,主要为x=8~16,既有多倍化也有非整倍化现象。染色体数据表明黄精族4个属的染色体进化模式各不相同,揭示了黄精族内染色体从高基数向低基数演化的规律;各属内染色体的演化主要是体现在二倍体水平上的核型变异,多倍化在本族中不占主导地位;仅黄精属内伴有非常强烈的非整倍化现象;细胞学证据与分子系统发育的结果比较吻合,为黄精族内属间以及属下的系统发育与进化提供了重要的参考资料。     

Genetic variability in Muscari comosum L. (Liliaceae) IV Geographical distribution and adaptive role of the polymorphic variants of chromosome 2

Muscari comosum L. (Liliaceae) has a chromosomal polymorphism for a pericentric inversion and a supernumerary chromosome segment probably due to an unequal interchange or insertional translocation. Both arrangements are widely distributed throughout the species range and the mean genetic distance among populations is D=0.131±0.075. There are no correlations between genetic distance and geographic distance or latitude. Only appreciable decreases in the frequencies of the inversion are detected in populations with ecologically marginal characteristics. There is a permanent and extended association between chromosomal inversion and an enzymatic locus (ADH). An excess of individuals heterozygous for the inversion was found and female productivity of heterozygotes is higher than that of corresponding homozygotes. A low rate of inversion heterozygosity in populations with ecologically marginal characteristics could be explained by natural selection. With respect to the adaptive role of the segment, although no homozygotes are found and may be selected against, heterozygotes could have heterotic effects.     

Monitoring of the genetic structure of natural populations: change of the effective population size and inversion polymorphism in <Emphasis Type="Italic">Drosophila subobscura</Emphasis>

We analyzed changes in the genetic structure and effective population size of two ecologically distinct populations of Drosophila subobscura over several years. Population sizes of D. subobscura in beech and oak wood habitats for a period of 6 years were estimated by the capture-mark-release-recapture method. Inversion polymorphism parameters were also assessed in the same populations for a period of 3 years. Significant differences in the numbers of individuals were observed between sexes. This affected the effective population sizes between particular years. The ratio of the effective size over the cenzus dropped significantly in beech wood in 2 years. Although overall heterozygosity remained unchanged during the years in both habitats, frequencies of gene arrangements on five chromosomes show variability. After the bottleneck, some complex chromosomal arrangements appeared for the first time in both populations. Standard gene arrangements of chromosome A increased in frequency over the years in each habitat, while the complex arrangements remain rather stable and specific for each population. The results obtained indicate that the population structure may significantly change if the effective size of D. subobscura population is reduced, which is mostly related to microclimatic changes in habitats. Based on the results to date, monitoring of microevolutionary changes by using D. subobscura and its relatives seems a promising way to study the effects of global climate changes.     

Inversion polymorphism in Southeast Asian populations of the Drosophila nasuta subgroup

Heterozygosity for chromosomal arrangements was investigated in four species of the nasuta complex. D. sulfurigaster albostrigata, D. albomicans and D. kohkoa are highly polymorphic whereas D. pulaua is monomorphic. Inversions were identified with the aid of photographic chromosome maps. The geographic distribution and frequencies of inversions detected and their possible phylogenies were discussed. Most inversions in D.s. albostrigata and D. albomicans occur on chromosome IIL. In D. kohkoa there is marked preponderance of inversions on chromosome III. Non-random association of certain inversions was noted. Variation in frequencies of inversions is interpreted as a result of adaptation to local ecogeographic conditions. Shared extant polymorphisms indicate phylogenetic relationships between species.The contents of this paper were incorporated in a Ph.D. thesis accepted by the University of Queensland in 1978. The author's present address is: Unit Genetik, Universiti Kebangsaan Malaysia, Jalan Pantai Baru, Kuala Lumpur, Malaysia.     

TRIVALENT FORMATION IN MULTIPLE TRISOMICS OF CLARKIA UNGUICULATA

Vasek, F. C. (U. California, Riverside.) Trivalent formation in multiple trisomics of Clarkia unguiculata. Amer. Jour. Bot. 50(3): 244–247. 1963.—A series of multiple trisomics, ranging from 2n + 1 to 2m + 7, was scored for univalents and trivalents at first metaphase. The mean number of trivalents per cell per extra chromosome was 0.43, 0.46, 0.46, 0.56, 0.54, 0.55 and 0.59 for plants with 1, 2, 3, 4, 5, 6 and 7 extra chromosomes respectively. These trivalent frequencies are interpreted to fall into 2 classes, indicating that the cytological differences between triploids and single trisomies may be based on a threshold effect. The results are compared with available data from other genera, and it is suggested that the observed increase in trivalent formation per extra chromosome may be associated with an increase in chiasma frequency.     

In situ hybridization (FISH) maps chromosomal homologies between Alouatta belzebul (Platyrrhini,Cebidae) and other primates and reveals extensive interchromosomal rearrangements between howler monkey genomes

We hybridized whole human chromosome specific probes to metaphases of the black-and-red howler monkey Alouatta belzebul in order to establish chromosomal homology between humans and black-and-red howlers. The results show that the black-and-red howler monkey has a highly rearranged genome and that the human chromosome homologs are often fragmented and translocated. The number of hybridization signals we obtained per haploid set was 40. Nine human chromosome probes gave multiple signals on different howler chromosomes, showing that their synteny is disturbed in A. Belzebul. Fourteen black-and-red howler autosomes were completely hybridized by one human autosomal paint, six had two signals, three had three signals, and one chromosome had four signals. Howler chromosomes with multiple signals have produced 12 chromosomal syntenies or hybridization associations which differ from those found in humans: 1/2, 2/20, 3/21, 4/15, 4/16, 5/7, 5/11, 8/18, 9/12, 10/16, 14/15, and 15/22. The hybridization pattern was then compared with those found in two red howler taxa and other mammals. The comparison shows that even within the genus Alouatta numerous interchromosomal rearrangements differentiate each taxa: A. belzebul has six unique apomorphic associations, A. seniculus sara and A. seniculus arctoidea share seven derived associations, and additionally A. seniculus sara has four apomorphic associations and A. seniculus arctoideaseven apomorphic associations. A. belzebul appears to have a more conserved karyotype than the red howlers. Both red and black-and-red howlers are characterized by Y-autosome translocations; the peculiar chromosomal sex system found in the red howler taxa could be considered a further transformation of the A. belzebul sex system. The finding that apparently morphologically similar or even identical taxa have such extreme genomic differences has important implications for speciation theory and neotropical primate conservation. Am. J. Primatol. 46:119–133, 1998. © 1998 Wiley-Liss, Inc.     

CYTOGEOGRAPHIC EVIDENCE FOR THE EVOLUTION OF DISTYLY FROM TRISTYLY IN THE NORTH AMERICAN SPECIES OF OXALIS SECTION IONOXALIS

A survey of haploid chromosome numbers of 18 North American taxa of Oxalis section Ionoxalis was initiated to determine the relationship between ploidal level, geographic distribution, and the occurrence of tristyly and distyly. Although chromosome numbers in the section are variable, the majority of tristylous populations are diploid. Among the distylous taxa a greater diversity of ploidal levels exists, with the higher chromosome numbers predominating. In section Ionoxalis the majority of the tristylous taxa are geographically restricted endemics of southern Mexico, while the distylous taxa have more extensive distributions ranging to the north. The association of diploidy and geographic endemism in the majority of the tristylous taxa suggests that these species are relictual. A few widespread tristylous taxa are polyploid, and often somewhat weedy. The probable derivation of widespread polyploid species from the restricted diploid endemics of southern Mexico appears to have been accompanied by the evolution of distyly from tristyly. The frequent association of polyploidy and distyly in section Ionoxalis has apparently resulted from the concurrence of two evolutionary trends: increase in ploidal level and the derivation of distyly from tristyly.     

Climate change and recent genetic flux in populations of <Emphasis Type="Italic">Drosophila robusta</Emphasis>

Background  

Studied since the early 1940's, chromosomal polymorphisms in the deciduous woods species Drosophila robusta have been characterized by well-defined latitudinal, longitudinal, and elevational clines, but – until at least ten years ago – stable, local population frequencies. Recent biogeographical analyses indicate that D. robusta invaded North America from southeast Asia and has persisted in eastern temperate forests for at least 20–25 my without speciating. The abundant chromosome polymorphisms found across the range of D. robusta are thus likely to be relatively ancient, having accumulated over many well known climatic cycles in North America. Sufficient long-term data are now available such that we can now gauge the rate of these evolutionary changes in natural populations due to environmental change.     

Selection in heterogeneous environments maintains the gene arrangement polymorphism of Drosophila pseudoobscura

Chromosomal rearrangements may play an important role in how populations adapt to a local environment. The gene arrangement polymorphism on the third chromosome of Drosophila pseudoobscura is a model system to help determine the role that inversions play in the evolution of this species. The gene arrangements are the likely target of strong selection because they form classical clines across diverse geographic habitats, they cycle in frequency over seasons, and they form stable equilibria in population cages. A numerical approach was developed to estimate the fitness sets for 15 gene arrangement karyotypes in six niches based on a model of selection-migration balance. Gene arrangement frequencies in the six different niches were able to reach a stable meta-population equilibrium that matched the observed gene arrangement frequencies when recursions used the estimated fitnesses with a variety of initial inversion frequencies. These analyses show that a complex pattern of selection is operating in the six niches to maintain the D. pseudoobscura gene arrangement polymorphism. Models of local adaptation predict that the new inversion mutations were able to invade populations because they held combinations of two to 13 local adaptation loci together.