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Allozyme variation and differentiation among four Apis species
Abstract
Summary — An electrophoretic survey of 18 putative enzyme loci was conducted on Sri Lankan Apis cerana, A. dorsata and A. florea, using techniques of sequential electrophoresis. Included for analysis were published data from European A. mellifera. Seven enzymes exhibited polymorphism and for some species are reported here for the first time. Seven enzymes showed fixed allozymic differences among all four species and three were fixed for the same allozyme in all four species. Allozyme data were analyzed using UPGMA clustering, a Distance Wagner procedure and a simple cladistic method. The mean genetic distance (Nei’s D = 1.30) indicates that considerable allozyme differences have accumulated among the species. Due to the relatively large genetic distance between A. cerana and A. mellifera (D = 1.10), electrophoretic data do not support a divergence time for these species as recent as that suggested by morphological and behavioral evidence. The electrophoretic data do not resolve questions concerning the phylogeny of Apis, due largely to the number of fixed differences among the species. Electrophoresis does show promise for species identification and resolution of the A. dorsata dcomplexd and other issues in Apis, as even the entry of data from single colonies into the BIOSYS program provided unambiguous discrimination. The treatment of electrophoretic data with phylogenetic tree-building or clustering techniques has potential for subspecies identification in endemic A. mellifera.
... Only small portions of the range of A. cerana have been sampled: Pakistan (Nunamaker et al., 1984), Sri Lanka (Sheppard and Berlocher, 1989), Thailand, peninsular Malaysia, southern Sulawesi and the Philippines (Gan et al., 1991), Yunnan, China (Li et al., 1986), Japan (Rozalski et al., 1996;Tanabe and Tamaki, 1985) and Korea (Lee and Woo, 1991;Lee et al., 1989). In addition sample sizes have been small, on the order of 13 or fewer colonies. ...
... Three studies (Gan et al., 1991;Lee and Woo, 1991;Sheppard and Berlocher, 1989) carried out a more thorough survey of 10-15 enzyme systems. Although a different suite of enzymes was surveyed in each study there is some overlap, particularly between the Korean (Lee and Woo, 1991) and Sri Lankan (Sheppard and Berlocher, 1989) studies. ...
... Three studies (Gan et al., 1991;Lee and Woo, 1991;Sheppard and Berlocher, 1989) carried out a more thorough survey of 10-15 enzyme systems. Although a different suite of enzymes was surveyed in each study there is some overlap, particularly between the Korean (Lee and Woo, 1991) and Sri Lankan (Sheppard and Berlocher, 1989) studies. Not surprisingly, studies that surveyed more enzyme systems detected more variation. ...
An analysis of the infraspecific categories of Apis cerana was prepared from the relevant literature on taxonomy, morphometrics, allozyme polymorphism and mtDNA diversity. About 31 puta- tive biometric groups have been proposed and assigned to about eight equivocal "subspecies" and var- ious ecotypes. However nearly half of the area of distribution of A. cerana remains unexamined. Allozyme polymorphism is greatest in southeast and lowest in northern and western Asia. About four major mtDNA groups are discernable. There is a very low overall geographic congruity amongst the morphoclusters, allozyme polymorphs and mtDNA clusters. The greatest problems in resolving infraspecific categories in A. cerana are inadequate sampling, incompatible differences in sample sizes, character suites, sampling distance, confidence limits and range of geographical scales employed in different studies.
... To-date, phylogenetic relationships between the species of the genus Apis are mainly based on morphological, ecoethological and biogeographic considerations (Ruttner, 1988). Recently, a phylogenetic tree has been obtained from allozyme data (Sheppard and Berlocher, 1989). In addition, preliminary reports of molecular systematics of Apis species have been presented by MacPheron (1989) and Sheppard (1989). ...
... The early separation of A florea, hypothetized by Michener (1974), is supported by a cladistic analysis of 19 morphological characters (Alexander, personal communication) and by the allozyme study of Sheppard and Berlocher (1989). Our sequence data, which are not opposed to this possi-bility, indicate that A florea and A dorsata diverged almost simultaneously of the cluster A mellifera -A cerana, as considered by Ruttner (1988, fig 3.11). ...
... To-date, phylogenetic relationships between the species of the genus Apis are mainly based on morphological, ecoethological and biogeographic considerations (Ruttner, 1988). Recently, a phylogenetic tree has been obtained from allozyme data (Sheppard and Berlocher, 1989). In addition, preliminary reports of molecular systematics of Apis species have been presented by MacPheron (1989) and Sheppard (1989). ...
... The early separation of A florea, hypothetized by Michener (1974), is supported by a cladistic analysis of 19 morphological characters (Alexander, personal communication) and by the allozyme study of Sheppard and Berlocher (1989). Our sequence data, which are not opposed to this possi-bility, indicate that A florea and A dorsata diverged almost simultaneously of the cluster A mellifera -A cerana, as considered by Ruttner (1988, fig 3.11). ...
The nucleotide sequence of the 5'end of the mitochondrial cytochrome-oxydase subunit 11 gene was obtained through direct sequencing of PCR (Polymerase Chain Reaction) product from 4 Apis species (mellifera, cerana, dorsata and florea) and Bombus lucorum (Apidae) and Xylocopa violacea (Anthophoridae) as outgroups. Phylogenetic trees were built using Neighbor-Joining and parsimony methods. Three branches, dorsata, florea and cerana-mellifera diverged almost simultaneously and cerana separated from mellifera later but not as early as generally thought.
... The sampling locations of the bees used here are listed in Supplemental Table S1. A. mellifera and A. cerana are known to be phylogenetically closer to each other than either is to A. dorsata (Alexander 1991; Arias and Sheppard 2005). Previous molecular dating suggested that A. mellifera and A. cerana were separated during the Miocene (6–8 million years ago [Mya]) (Sheppard and Berlocher 1989; Garnery et al. 1991). We first targeted region 1 of the gene (Fig. 1) because insertions and deletions, which may lower the reliability of phylogenetic analyses, are less frequent in this region than in other regions of the gene, especially region 3. ...
... It is interesting to determine the age of the csd alleles. The average number of nucleotide substitutions per site (d) between A. mellifera and A. cerana is 0.07 in the six neutral genomic regions sequenced (Table 1), and the two species diverged ∼7 Mya (Sheppard and Berlocher 1989; Garnery et al. 1991 ). Therefore , the honey bee molecular clock ticks at a rate of ∼10 substitutions per kilobase per million years, which is close to that in fruit flies (11.1) (Tamura et al. 2004). ...
The mechanism of sex determination varies substantively among evolutionary lineages. One important mode of genetic sex determination is haplodiploidy, which is used by approximately 20% of all animal species, including >200,000 species of the entire insect order Hymenoptera. In the honey bee Apis mellifera, a hymenopteran model organism, females are heterozygous at the csd (complementary sex determination) locus, whereas males are hemizygous (from unfertilized eggs). Fertilized homozygotes develop into sterile males that are eaten before maturity. Because homozygotes have zero fitness and because common alleles are more likely than rare ones to form homozygotes, csd should be subject to strong overdominant selection and negative frequency-dependent selection. Under these selective forces, together known as balancing selection, csd is expected to exhibit a high degree of intraspecific polymorphism, with long-lived alleles that may be even older than the species. Here we sequence the csd genes as well as randomly selected neutral genomic regions from individuals of three closely related species, A. mellifera, Apis cerana, and Apis dorsata. The polymorphic level is approximately seven times higher in csd than in the neutral regions. Gene genealogies reveal trans-species polymorphisms at csd but not at any neutral regions. Consistent with the prediction of rare-allele advantage, nonsynonymous mutations are found to be positively selected in csd only in early stages after their appearances. Surprisingly, three different hypervariable repetitive regions in csd are present in the three species, suggesting variable mechanisms underlying allelic specificities. Our results provide a definitive demonstration of balancing selection acting at the honey bee csd gene, offer insights into the molecular determinants of csd allelic specificities, and help avoid homozygosity in bee breeding.
... In small populations or in those having experienced a bottleneck , the overall coalescence times are expected to be shorter and the average sequence divergence lower because alleles are more likely to be lost by genetic drift. In this study, we analyze csd gene sequences of 3 related honeybee species, Apis cerana, Apis dorsata (both Asian honeybee species), and A. mellifera (the western honeybee ) that diverged over the last 10 Myr (Sheppard and Berlocher 1989; Garnery et al. 1992, see Materials and Methods). These species share very similar life histories, have a highly social organization, and show substantial division of labor and reproduction (i.e., they are eusocial). ...
... An estimate of the genomic mutation rate (7 Â 10 À9 per site per year) is derived from the average pairwise synonymous divergence per site (dS 5 0.1) of fem and elongation factor-alpha 1 (EF-a 1) sequences of A. mellifera and A. cerana and estimates of their divergence times (;7 Myr) (Sheppard and Berlocher 1989; Garnery et al. 1992) according to the following relationship: number of polymorphisms at synonymous sites that accumulate 52gl, where g is the number of generations, assuming 1 generation per year for the honeybee, and l is the mutation rate. From the mutation rate and average synonymous divergence between A. mellifera and A. dorsata for fem and EF-a 1 sequences (dS 5 0.135), a rough estimate of their divergence is obtained (10 Myr). ...
Our understanding of the impact of recombination, mutation, genetic drift, and selection on the evolution of a single gene is still limited. Here we investigate the impact of all these evolutionary forces at the complementary sex determiner (csd) gene that evolves under a balancing mode of selection. Females are heterozygous at the csd gene and males are hemizygous; diploid males are lethal and occur when csd is homozygous. Rare alleles thus have a selective advantage, are seldom lost by the effect of genetic drift, and are maintained over extended periods of time when compared with neutral polymorphisms. Here, we report on the analysis of 17, 19, and 15 csd alleles of Apis cerana, Apis dorsata, and Apis mellifera honeybees, respectively. We observed great heterogeneity of synonymous (piS) and nonsynonymous (piN) polymorphisms across the gene, with a consistent peak in exons 6 and 7. We propose that exons 6 and 7 encode the potential specifying domain (csd-PSD) that has accumulated elevated nucleotide polymorphisms over time by balancing selection. We observed no direct evidence that balancing selection favors the accumulation of nonsynonymous changes at csd-PSD (piN/piS ratios are all <1, ranging from 0.6 to 0.95). We observed an excess of shared nonsynonymous changes, which suggest that strong evolutionary constraints are operating at csd-PSD resulting in the independent accumulation of the same nonsynonymous changes in different alleles across species (convergent evolution). Analysis of csd-PSD genealogy revealed relatively short average coalescence times ( approximately 6 Myr), low average synonymous nucleotide diversity (piS < 0.09), and a lack of trans-specific alleles that substantially contrasts with previously analyzed loci under strong balancing selection. We excluded the possibility of a burst of diversification after population bottlenecking and intragenic recombination as explanatory factors, leaving high turnover rates as the explanation for this observation. By comparing observed allele richness and average coalescence times with a simplified model of csd-coalescence, we found that small long-term population sizes (i.e., N(e) < 10(4)), but not high mutation rates, can explain short maintenance times, implicating a strong historical impact of genetic drift on the molecular evolution of highly social honeybees.
... This analysis allows the study of the origin of honey bees [88,121]. Furthermore, some studies based on allozyme and mtDNA variations support the findings of Ruttner (1988) [72,99,[122][123][124][125]. ...
Biological diversity of bee - the genetic resource in Russia, which enables maintaining homeostasis of ecosystems through pollination entomophilies plants. The biodiversity of bees in human life has ecological, social, economic and aesthetic significance. Of particular interest in the preservation of biodiversity are taxonomically isolated species and populations, not resemble others and therefore unique in their genetic constitution. These species are often endemic, that is limited to the dissemination of one area. Their extinction of will mean the loss of biodiversity. Uncontrolled introduction of bees of different species breeds and populations leads to the spread of diseases and hidden genetic defects. In the process of mass introduction of not adapted breeds of honey bees there is a loss of breed diversity of endemic populations, accompanied by a narrowing of breeding opportunities and a reduction in pollinators. Using modern methods of monitoring with the use of microsatellite analysis to improve the efficiency study of the gene pool of honeybees. Biotechnological methods of artificial insemination of Queens and cryopreservation of drone sperm in liquid nitrogen allow preserving the gene pool of endangered native breeds of honeybees. The use of these methods makes it possible to avoid polyandry and conduct controlled mating in breeding and genetic studies. Obtaining a culture of honeybee cells is promising for a more in-depth study of the interaction with intracellular infectious agents, genomic and epigenetic mechanisms of variability of this unique object.
... This analysis allows the study of the origin of honey bees [88,121]. Furthermore, some studies based on allozyme and mtDNA variations support the findings of Ruttner (1988) [72,99,[122][123][124][125]. ...
The loss of honey bees has drawn a large amount of attention in various countries. Therefore, the development of efficient methods for recovering honey bee populations has been a priority for beekeepers. Here we present an extended literature review and report on personal communications relating to the characterization of the local and bred stock of honey bees in the Russian Federation. New types have been bred from local colonies (A. mellifera L., A. m. carpatica Avet., A. m. caucasia Gorb.). The main selection traits consist of a strong ability for overwintering, disease resistance and different aptitudes for nectar collection in low and high blooming seasons. These honey bees were certified by several methods: behavioral, morphometric and genetic analysis. We illustrate the practical experience of scientists, beekeepers and breeders in breeding A. mellifera Far East honey bees with Varroa and tracheal mite resistance, which were the initial reasons for breeding the A. mellifera Far Eastern breed by Russian breeders, Russian honey bee in America, the hybrid honey bee in Canada by American breeders, and in China by Chinese beekeepers. The recent achievements of Russian beekeepers may lead to the recovery of beekeeping areas suffering from crossbreeding and losses of honey bee colonies.
... Other honey bee remains are only known from archaeological sites, at most several thousand years old 91,92 . As such, studies have so far yielded highly divergent results; the split between A. cerana and A. mellifera has been variously estimated as having occurred between 6 and 25 million years ago 38,72,93,94 and the rapid divergence between A. mellifera subspecies between 0.3 and 1.3 million years ago 6,38,75 . Wallberg et al. 39 used the genealogical concordance method to estimate that honey bee subspecies diverged between 13,000-38,000 years ago, which would correspond to the last glacial maximum, implying that the expansion of A. mellifera from its region of origin into Europe began after the retreat of the ice sheets. ...
Western honey bees (Apis mellifera) are one of the most important pollinators of agricultural crops and wild plants. Despite the growth in the availability of sequence data for honey bees, the phylogeny of the species remains a subject of controversy. Most notably, the geographic origin of honey bees is uncertain, as are the relationships among its constituent lineages and subspecies. We aim to infer the evolutionary and biogeographical history of the honey bee from mitochondrial genomes. Here we analyse the full mitochondrial genomes of 18 A. mellifera subspecies, belonging to all major lineages, using a range of gene sampling strategies and inference models to identify factors that may have contributed to the recovery of incongruent results in previous studies. our analyses support a northern African or Middle eastern origin of A. mellifera. We show that the previously suggested european and Afrotropical cradles of honey bees are the result of phylogenetic error. Monophyly of the M, c, and o lineages is strongly supported, but the A lineage appears paraphyletic. A. mellifera colonised europe through at least two pathways, across the Strait of Gibraltar and via Asia Minor.
... DNA sequencing of intergenic region between tRNA leu region and the COII gene of honey bees A. mellifera, A. cerana, A. dorasata, A. andreniformis and A. florea revealed that there was a 7-11% sequence divergence between these five Apis species. Cornuet and Garney (1991) presented a phylogeny tree developed from the mtDNA sequence, which showed early divergence of A. florea with close affinity towards cavity nesting bees viz., A. mellifera and A.cerana which was in agreement with morphometrical, behavioural (Alexander, 1991) and allozyme data (Sheppard and Berlocher, 1989). Contrary to the above, the phylogenetic tree of six Apis species including two dwarf honey bees constructed using COII gene sequence has revealed that A. dorasata was the most ancestral group. ...
... The Asian honey bee (Apis dorsata) is an important pollinator and source of honey throughout its natural range in Southeast and East Asia (Chantawannakul 2018;Sheppard and Berlocher 1989;Thapa 2006). This species is distinct from the European honey bee (Apis mellifera) both morphologically and behaviorally. ...
The Asian honey bee (Apis dorsata) is distinct from its more widely distributed cousin A. mellifera by a few key characteristics. Most prominently, A. dorsata, nest in the open by forming a colony clustered around the honeycomb, while A. mellifera nest in concealed cavities. Additionally, the worker and reproductive castes are all of the same size in A. dorsata. In order to investigate these differences, we performed whole genome sequencing of A. dorsata using a hybrid Oxford Nanopore and Illumina approach. The 223MB genome has an N50 of 35kb with the largest scaffold of 302kb. We have found that there are many genes in the dorsata genome that are distinct from other hymenoptera and also large amounts of transposable elements, and we suggest some candidate genes for A. dorsata's exceptional level of defensive aggression.
... The Asian honey bee (Apis dorsata) is an important pollinator and source of honey throughout its natural range in Southeast and East Asia [1][2][3]. This species is distinct from the European honey bee (Apis mellifera) both morphologically and behaviorally. ...
The Asian honey bee (Apis dorsata) is distinct from its more widely distributed cousin A. mellifera by a few key characteristics. Most prominently, A. dorsata, nest in the open by forming a colony clustered around the honeycomb, while A. mellifera nest in concealed cavities. Additionally, the worker and reproductive castes are all of the same size in A. dorsata. In order to investigate these differences, we performed whole genome sequencing of A. dorsata using a hybrid Oxford Nanopore and Illumina approach. The 223MB genome has an N50 of 35kb with the largest scaffold of 302kb. We have found that there are many genes in the dorsata genome that are distinct from other hymenoptera and also large amounts of transposable elements, and we suggest some candidate genes for A. dorsata's exceptional level of defensive aggression.
... Ces chiffres ne prennent pas en compte le nombre inquantifiable d'apiculteurs qui viennent transhumer dans le PnC et qui déclarent leur activité dans la commune de leur siège d'exploitation située hors du Parc. Garnery et al. 1991, Sheppard & Berlocher 1989, Engel 1999, Arias & Sheppard 1996, 2005)Engel 1999, Sheppard & Meixner, 2003, Rortais et al. 2010)Cévennes, cette description est partagée dans le discours des personnes âgées lorsqu'elles évoquent leurs souvenirs de la première moitié du XX e siècle. L'abeille qui peuple alors les ruches-troncs, est l'abeille locale, mais il n'est pas souvent question de l'abeille noire à proprement parler. ...
On the grounds that beekeeping of the single domesticated honeybee looks at first sight homogenous throughout whole Europe, understanding the evolution of this activity at the very local level is generally overlooked. Such understanding is nonetheless crucial to conceive appropriate community-based resource management in socioecological systems that have been profoundly imprinted by beekeeping activities. We carried out an ethnoecological history study focusing on local knowledge regarding beekeeping in the Cevennes National Park (southern France). By combining the analysis of archival documents, scientific literature and personal testimonies from elderly Cevennes dwellers, this dive into history has allowed us to reconstitute the major episodes of beekeeping in Cevennes, by considering the modifications of chosen beehive models and bee landraces, as well as the valorization of beehive products following the evolving social and economic circumstances. Most salient features, such as the first evidence of the use of log hives in the early 17th century, were thus time-stamped and retrospectively set into context. Artisanal beekeeping of the local black bee hosted in log hives has persisted until the 1970s date of the transition to modern beekeeping using frame hives, selected bee landraces, and a professionalization of the local honey trade sector. Beekeepers from the Cevennes region only lately stepped from a domestic and landscaped beekeeping, which was optimized in a context of self-sufficient pluriactivity, into an intensive beekeeping driven by the search for maximized honey yields and supported by a diversification and a hybridization of bee landraces. Such combined historical and biocultural perspectives of beekeeping in Cevennes should serve to elaborate reasonable goals for conservation and should help conciliating the preservation of a patrimonial and traditional beekeeping along with the enhancement of a yet emerging local honeybee market.
... Allozymes are markers exhibiting only few alleles in honey bees and other Hymenoptera as consequence of male haploidy (Pamilo and Crozier, 1981). Within A. mellifera, no fixed allelic differences between populations exist; in consequence, variation between populations consists exclusively of differences in allele frequencies, with a skewed distribution of alleles and mostly one very dominant allele (Sheppard and Berlocher, 1989). Although differences in allele frequencies have been successfully used to separate honey bee subspecies, this differentiation resulted mainly from rare or private alleles (Cornuet, 1986). ...
The COLOSS GEI (Genotype-Environment Interactions) Experiment was setup to further our understanding of recent honey bee colony losses. The main objective of the GEI experiment was to understand the effects of environmental factors on the vitality of European honey bee genotypes. This paper aims to describe the genetic background and population allocation of the bees used in this experiment. Two wing morphometric and two genetic methods were employed to discriminate bee populations. Classical morphometry of 11 angles on the wings were carried out on 350 bees. Geometric morphometry on 19 wing landmarks was carried out on 381 individuals. DNA microsatellite analysis was carried out on 315 individuals using 24 loci. Allozyme analysis was performed on 90 individuals using six enzyme systems. DNA microsatellite markers produced the best discrimination between the subspecies (Apis mellifera carnica, A. m. ligustica, A. m. macedonica, A. m. mellifera and A. m. siciliana) used in the experiment. Morphometric methods generally showed an intermediate level of discrimination, usually best separating A. m. siciliana and A. m. ligustica from the remaining populations. Allozyme markers lack power to discriminate at the level of individual bees, and given our sample size, also fail to differentiate subspecies. Based on DNA microsatellites, about 69% of the individuals were assigned to the same subspecies as originally declared, and 17% were found to belong to a different subspecies. Fourteen percent of the samples were found to be of mixed origin and could not be assigned to any subspecies with certainty. We further discuss the caveats of the methods and details of the sampled bees, their origins and breeding programmes in their respective locations.
... However, the foragers in that study were still able to locate feeders in the vicinity of the food source, even after having followed dances in a different dialect. While the subspecies of Apis mellifera may have diverged around 0.67 million years ago [30], [31], our study confirms that the ability to use the information encoded in an unfamiliar dance extends even across species separated by six to eight million years of evolution [32], [33]. The Acc bees in our mixed-species colony were almost as successful as the Aml bees in locating a feeder advertised by more experienced Aml dancers, as indicated by our accuracy experiments. ...
The honeybee waggle dance, through which foragers advertise the existence and location of a food source to their hive mates, is acknowledged as the only known form of symbolic communication in an invertebrate. However, the suggestion, that different species of honeybee might possess distinct 'dialects' of the waggle dance, remains controversial. Furthermore, it remains unclear whether different species of honeybee can learn from and communicate with each other. This study reports experiments using a mixed-species colony that is composed of the Asiatic bee Apis cerana cerana (Acc), and the European bee Apis mellifera ligustica (Aml). Using video recordings made at an observation hive, we first confirm that Acc and Aml have significantly different dance dialects, even when made to forage in identical environments. When reared in the same colony, these two species are able to communicate with each other: Acc foragers could decode the dances of Aml to successfully locate an indicated food source. We believe that this is the first report of successful symbolic communication between two honeybee species; our study hints at the possibility of social learning between the two honeybee species, and at the existence of a learning component in the honeybee dance language.
... Biochemical variation among the highly distinctive bee species, A cerana, A dorsata and A florea (Francis et al, 1985;Sheppard and Berlocher, 1989) is apparently greater than that detected here among the A dorsata group. The biochemical variation detected within our A dorsata collection is similar to that seen among subspecies of A mellifera (Sylvester, 1986;Sheppard and Huettel, 1988). ...
Cuticular hydrocarbon pattern (CHP) analysis was performed on giant honey bees (the Apis dorsata group) including: 1), those occasionally given species status-Himalayan honey bees, Philippine honey bees, Sulawesi honey bees; 2), those separated since the Pleistocene-common A dorsata of the Indian and Asian lowlands and islands on the continential shelf (India and Sri Lanka, Thailand and Sumatra); and 3), giant honey bees of Borneo and Palawan, potential stepping-stones to the Phillippines and Sulawesi. Four groups were found among giant honey bees by this CHP analysis. Most distinctive were those of Palawan and Nepal. The widespread lowland Apis dorsata differed very little among mainland and island populations, whereas those of Borneo, Sulawesi, and Phillippines proper formed a single group. Those of the Himalayas appear to have diverged from A dorsata.
... However, the foragers in that study were still able to locate feeders in the vicinity of the food source, even after having followed dances in a different dialect. While the subspecies of Apis mellifera may have diverged around 0.67 million years ago [30], [31], our study confirms that the ability to use the information encoded in an unfamiliar dance extends even across species separated by six to eight million years of evolution [32], [33]. The Acc bees in our mixed-species colony were almost as successful as the Aml bees in locating a feeder advertised by more experienced Aml dancers, as indicated by our accuracy experiments. ...
The honeybee waggle dance, through which foragers advertise the existence and location of a food source to their hive mates, is acknowledged as the only known form of symbolic communication in an invertebrate. However, the suggestion, that different species of honeybee might possess distinct 'dialects' of the waggle dance, remains controversial. Furthermore, it remains unclear whether different species of honeybee can learn from and communicate with each other. This study reports experiments using a mixed-species colony that is composed of the Asiatic bee Apis cerana cerana (Acc), and the European bee Apis mellifera ligustica (Aml). Using video recordings made at an observation hive, we first confirm that Acc and Aml have significantly different dance dialects, even when made to forage in identical environments. When reared in the same colony, these two species are able to communicate with each other: Acc foragers could decode the dances of Aml to successfully locate an indicated food source. We believe that this is the first report of successful symbolic communication between two honeybee species; our study hints at the possibility of social learning between the two honeybee species, and at the existence of a learning component in the honeybee dance language.
... While there are no inconsistencies among these three studies , available information is insufficient to apply to the whole area of A. florea distribution. Similarly, available data on enzyme polymorphism in A. florea (Li et al., 1986; Sheppard and Berlocher, 1989; Gan et al., 1991) are likewise geographically limited precluding extrapolation to the whole A. florea population. The available genetic data are too regional in nature to be informative for the species as a whole. ...
Multivariate morphometric analyses were performed on 2923 individual worker bees from 184 colonies representing 103 localities across the full distributional area of Apis florea Fabricius 1787 from Vietnam and southeastern China to Iran and Oman (~7000 km). Morphologically A. florea is unequivocally separable from A. andreniformis. Comparisons of geographically separated A. florea populations result in morphoclusters that reflect sampling artifacts. These morphoclusters change clinally with latitude but overlap when the full database is contained in the same principal component analysis. A cluster analysis based on Euclidean distances suggests degrees of affinity between various geographic groupings of A. florea. This species occupies a large area that includes rainforests, savannas, subtropical steppes, and semideserts. The seasonality of reproductive swarming is temporally continuous allowing gene flow throughout this panmictic species.
... While the relative youth of a million-yearold A. mellifera lineage has been supported by molecular studies, cladogenesis of A. mellifera and A. cerana appears to have occurred at a much earlier time (6-9 mya) based on allozyme and DNA sequence differences (Sheppard and Berlocher, 1989;Cornuet and Garnery, 1991;. The apparent discrepancy between the age of A. mellifera subspecies and the A. mellifera/A. ...
Endemic honey bees of the Tien Shan Mountains in Central Asia are described as a new subspecies, Apis mellifera pomonella, on the basis of morphometric analyses. Principal component and discriminant analysis of the morphological characters measured clearly place these bees into the oriental evolutionary branch of honey bees, but also show that they are distinct from the other subspecies in this lineage. The existence of this newly described honey bee subspecies extends the range of endemic A. mellifera more than 2000 km eastward than previously estimated. Sequence analysis of mitochondrial DNA places A. m. pomonella within the C mitochondrial lineage (a group that is inclusive of both C and O morphological lineages). These findings support the conclusion that A. m. pomonella has a phylogeographic history shared with subspecies from the eastern limit of the previously known range.
... Genetic diversity of managed stocks of honey bees, Apis mellifera L., has been measured throughout the United States and Italy (Sheppard and Berlocher 1989, Schiff and Sheppard 1995, Schiff and Sheppard 1996, Bourgeois et al. 2008. Maintenance of genetic diversity among breeding lines is an important factor in selective breeding and stock management. ...
Maintenance of genetic diversity among breeding lines is important in selective breeding and stock management. The Russian Honey Bee Breeding Program has strived to maintain high levels of heterozygosity among its breeding lines since its inception in 1997. After numerous rounds of selection for resistance to tracheal and varroa mites and improved honey production, 18 lines were selected as the core of the program. These lines were grouped into three breeding blocks that were crossbred to improve overall heterozygosity levels of the population. Microsatellite DNA data demonstrated that the program has been successful. Heterozygosity and allelic richness values are high and there are no indications of inbreeding among the three blocks. There were significant levels of genetic structure measured among the three blocks. Block C was genetically distinct from both blocks A and B (F(ST) = 0.0238), whereas blocks A and B did not differ from each other (F(ST) = 0.0074). The same pattern was seen for genic (based on numbers of alleles) differentiation. Genetic distance, as measured by chord distance, indicates that all of the 18 lines are equally distant, with minimal clustering. The data indicate that the overall design of the breeding program has been successful in maintaining high levels of diversity and avoiding problems associated with inbreeding.
... In insects, esterases catalyze the hydrolysis of ester bonds and have been shown to participate in digestive processes, in the regulation of juvenile hormone production and in the degradation of insecticides. Sheppard and Berlocher (1989) used enzymatic systems to demonstrate the existence of inter-specific and intra-specific variation in four species of Apis. Scarpassa et al. (1996) studied the profile of esterases in six populations of Anopheles nuneztovari and observed that the enzymes at locus EST5 had high levels of allelic variability among Brazilian populations but low variability among Colombian populations. ...
This report describes a case of hybridization between two species of Meliponinae bees (Melipona scutellaris from the Diamantina plateau in the State of Bahia, and Melipona capixaba from around Domingos Martins, in the State of Espírito Santo, Brazil). To demonstrate hybridization, electrophoretic profiles of esterase activity from three colonies were studied. Ten adult workers of M. scutellaris, M. capixaba and the hybrid colony were collected and processed individually. The pattern of esterase activity was constant for each species but differed between them, whereas hybrid bees had a pattern derived from both species. The fact that two ecologically different species of stingless bees separated by more than 300 km could still cross when placed in the same area suggests that there has not been any pressure to develop reproductive isolation.
... As indicated in the results section, the MDH-1 * , EST-3 * , and SOD * loci were polymorphic in all the populations studied. By comparing our results with those of similar studies, it should be emphasized that no SOD * locus polymorphism had been found for A. mellifera subspecies (Sheppard and Berlocher, 1989). The usefulness of the MDH-1 * and EST-3 * loci, as reported by Sheppard and Smith (2000), is strengthened by our data (three alleles for MDH-1 * and two alleles for EST-3 * ). ...
Ten gene enzymic systems (α-GPDH, AO, MDH, ADH, LAP, SOD, ALP, ACPH, ME, and EST), corresponding to 12 genetic loci, were assayed from five Greek populations representing three subspecies of Apis mellifera, A. m. cecropia (Pthiotida, Kythira), A. m. macedonica (Macedonia), and the “Aegean race” of A. mellifera, which is supposed to be very similar to A. m. adami (Ikaria, Kasos), as well as a population from Cypus (A. m. cypria). ADH
∗-1, ADH
∗-2, and LAP
∗ electrophoretic patterns discriminate the Cyprus population from the Greek populations. MDH
∗-1, EST
∗-3, SOD
∗, ALP
∗, and ME
∗ loci were found to be polymorphic in almost all populations. The observed heterozygosity was found to range from 0.066 to 0.251. Allele frequencies of all loci were used to estimate Nei's genetic distance, which was found to range between 0.011 and 0.413 among the populations studied. UPGMA and neighbor-joining phylogenetic trees obtained by genetic distance matrix methods, as well as a Wagner tree based on the discrete character parsimony method, support the hypothesis that the most distant population is that from Cyprus. Our allozymic data support A. m. cypria as a distinct subspecies, but there was no allozymic support for the distinction of the other subspecies existing in Greece.
Arthropods of kingdom animalia are bountiful on earth and copious of biodiversity is yet to be discovered. Traditional taxonomy identifies the species based on external morphological, anatomical, and ecological characters is tedious with few of limitations for describing the species. While DNA barcoding became an alternative and quick tool to delineate the species and it has been practiced by many researchers across the world. The DNA barcoding refers to sequencing of a short fragment of mitochondrial cytochrome c oxidase subunit I (COX I) to identify species, including unknown species. Apart from systematics, it also found to have a role in biological control, bio-surveillance, and quality control in food industry, conservation of endangered species and integrated pest management. However, few researchers are criticizing this method due to nuclear mitochondrial DNA (NUMTs) and mitochondrial DNA introgression between closely related species facilitated by Wolbachia, which will lead to over-estimation of species.
The thesis for the degree of candidate of biological sciences. Preservation of the gene pool of dark forest honey bee Ammellifera is the base point for the development of beekeeping and bee breeding in Russia
which in turn requires solving a number of problems in basic and
applied research. Our work touches the three of them. We were able to prove
the existence of the Urals several genetic reserves that should be the basis for the preservation of the gene pool Ammellifera. It was expanded set of genetic markers and statistical approaches used for this purpose. We
It managed to obtain new data on the structure of intra honeybee
A.mellifera. We hope that the results will allow to approach
the problem of preserving the gene pool Ammellifera both in Russia and in other Nordic countries.
As is known, the use morphometric methods for this purpose little information when using hybrid bees. It is shown that by using
molecular methods may determine which subspecies belong to the bees.
However, not all genetic loci can become markers. Only
strict selection and integrated use of nuclear loci and
mitochondrial genomes will be effective in the identification of subtypes of bees. Equally important in the study played a statistical analysis of
results. Statistical methods, selection of genetic markers and the character of the collection object is strictly dependent on the goals of the study.
Phylogenetic analysis also allowed for the first time to establish close
genetic kinship bees Ural and European populations Ammellifera.
In parallel, we have been able to show that the evolutionary branch of M consists only of subspecies Ammellifera, whereas previously it was thought that it includes several other
A.mellifera subspecies of the North-West Africa and the Iberian Peninsula.
The obtained genetic characteristics of populations of bees Urals
unique and will serve as a basis for further research, as well as
measures for the conservation and restoration of the entire gene pool Ammellifera
the territory of the former habitat subspecies. Furthermore, based on these data, we can
develop a strategy for the further development and management of beekeeping
Ammellifera populations in Russia and Europe.
On the basis of polymorphism intergenic locus COI-COII mitochondrial
DNA demonstrated the existence of at least four stored locally
populations of Apis mellifera mellifera L. in the Urals: Vishera, Kama region of the South,
tatyshlinskoy and Burzyan.
An analysis of the frequency of occurrence of combinations of PQQ intergenic locus COI-COII
mtDNA showed that uinskaya population, previously hosted by morphometric
featured for the population Ammellifera, it is a hybrid. Genetic distance
Ammellifera between the Ural and the hybrid population can be Iglinsky
accepted for further studies at one of the criteria for the safety of aboriginal
population A.m.mellifera.
A study of polymorphism loci separate nuclear and mitochondrial
DNA revealed a close genetic relationship, a small fraction of inbreeding and a lack of
heterozygotes in the Ural population Ammellifera.
Phylogenetic analysis based on a comparison of the nucleotide
ND2 gene fragment sequences of mitochondrial DNA revealed
genetic kinship of the Ural and Western European populations Ammellifera.
Comparative analysis mitotypes gene fragment of the mitochondrial ND2
DNA shows that subspecies Ammellifera, probably the only
Representative intraspecific evolutionary branches of M, which is thus not
It should include not only the African subspecies Apis mellifera sahariensis
Baldensperger and Apis mellifera intermissa Maa, but Spanish subspecies of Apis mellifera
iberica Goetze. It found that species ancestral form Apis mellifera L., could be
With bee evolutionary branch, and not, as previously thought.
Диссертация на соискание ученой степени кандидата биологических наук. Сохранение генофонда тёмной лесной медоносной пчелы A.m.mellifera являлось базовым моментом для развития пчеловодства и селекции пчел в России,
что в свою очередь требует решения целого ряда задач в области фундаментальных и
прикладных исследований. Наша работа касалась трёх из них. Нам удалось доказать
существование на Урале нескольких генетических резерватов, которые должны стать основой для сохранения генофонда A.m.mellifera. Был расширен комплекс генетических маркёров и статистических подходов, используемых для этого. Нам
удалось получить новые данные по внутривидовой структуре медоносной пчелы
A.mellifera. Мы надеемся, что полученные результаты позволят приблизиться к
решению проблемы сохранения генофонда A.m.mellifera как в России, так и в других странах Северной Европы.
Как известно, использование морфометрических методов для этой цели малоинформативно при работе с гибридными пчелами. Показано, что с помощью
молекулярных методов возможно определить, к какому подвиду принадлежат пчелы.
Тем не менее, не любые локусы могут стать генетическими маркерами. Только
строгий подбор и комплексное использование локусов ядерного и
митохондриального геномов будет эффективно в идентификации подвидов пчел. Не менее важную роль в исследовании играла статистическая обработка полученных
результатов. Методы статистической обработки, подбор генетических маркеров и характер сбора объекта строго зависят от поставленных целей исследования.
Также филогенетический анализ позволил впервые установить тесное
генетическое родство пчел уральских и европейских популяций A.m.mellifera.
Параллельно мы смогли показать, что эволюционная ветвь М состоит только из подвида A.m.mellifera, тогда как раньше считалось, что в нее входят несколько других
подвидов A.mellifera из Северо-Западной Африки и Иберийского полуострова.
Полученные нами генетические характеристики популяций пчел Урала
уникальны и будут служить основой для дальнейших исследований, а также
мероприятий по сохранению и восстановлению генофонда A.m.mellifera на всей
территории прежнего ареала подвида. Кроме того, на основе этих данных мы можем
разработать стратегию дальнейшего развития пчеловодства и управления
популяциями A.m.mellifera в России и Европе.
На основе полиморфизма межгенного локуса COI-COII митохондриальной
ДНК показано существование как минимум четырёх сохранившихся локальных
популяций Apis mellifera mellifera L. на Урале: вишерской, южно-прикамской,
татышлинской и бурзянской.
Анализ частот встречаемости комбинации PQQ межгенного локуса COI-COII
мтДНК показал, что уинская популяция, ранее принимаемая по морфометрическим
признакам за популяцию A.m.mellifera, являлась гибридной. Генетическое расстояние
между уральской A.m.mellifera и гибридной иглинской популяцией может быть
принято в дальнейших исследованиях за один из критериев сохранности аборигенной
популяции A.m.mellifera.
Исследование полиморфизма отдельных локусов ядерной и митохондриальной
ДНК выявило тесное генетическое родство, небольшую долю инбридинга и дефицит
гетерозигот в уральских популяциях A.m.mellifera.
Филогенетический анализ на основе сравнения нуклеотидных
последовательностей фрагмента гена ND2 митохондриальной ДНК показал
генетическое родство уральских и западноевропейских популяций A.m.mellifera.
Сравнительный анализ митотипов фрагмента гена ND2 митохондриальной
ДНК свидетельствует, что подвид A.m.mellifera, вероятно, является единственным
представителем внутривидовой эволюционной ветви М, в которую, таким образом, не
следует включать не только африканские подвиды Apis mellifera sahariensis
Baldensperger и Apis mellifera intermissa Maa, но и испанский подвид Apis mellifera
iberica Goetze. Обнаружено, что предковой формой вида Apis mellifera L., могли быть
пчелы эволюционной ветви С, а не О, как считалось ранее.
Ilyasov R.A. Polymorphism of Apis mellifera mellifera L. in Ural. Thesis of Ph.D. 2006. 204 p.
Samples of Apis mellifera ligustica from Emilia (northcentral Italy), samples of A. m. carnica from Austria and from northern Yugoslavia, and samples from Friuli (northeast Italy), regarded as a zone of hybridization, were assayed for allozyme polymorphism. Five loci were found to be polymorphic: Est-3, Est6, Mdh-l, Me, Pgm. Est-6 was used for the first time in a wide population study of A. mellifera. Among the polymorphic loci reported to date, Est-6 seems to be the more distinctive between A. m. ligustica and A. m. carnica. By a distance Wagner analysis of electrophoretic data, the samples of the two races were separated into two clearly distinct phenogram clusters. Friuli samples were clustered in an intermediate position.
This chapter focuses on the molecular biology of the honeybee. The biological rules that govern the honeybee colony have fascinated scientists since Aristotle. The honeybee riddle of chaos on the one hand and yet pattern on the other hand was to occupy generations of scientists after Aristotle. The focus on behavioral ecology was not always so strong, and in fact honeybees also entered the arena of modern research as a genetic model system. The molecular revolution of genetics took place leaving the honeybees aside. The molecular genetics of Drosophila melanogaster boomed, while bee geneticists felt more attracted to applied breeding research. Because honeybees are of significant economical and ecological importance, there is obviously also a compelling need to understand their population and breeding genetics. The molecular genetics of honeybees can both increase the understanding of basic genetic mechanisms and improve the knowledge of honeybee specific genetic problems. The mitochondria1 genome of the honeybee is much better known than its nuclear counterpart. The segmentation of the embryo is one of the best understood steps in insect embryogenesis. In particular, the findings in Drosophila melanogaster have given an important insight into gene regulation in early embryonic development. The pattern of gene activity corresponds to the external metameric subdivision in the larval body. Genetic variability in populations has been analyzed with isozyme polymorphisms. Although this technique has proved very powerful for the study of many insect populations, isozyme analysis in honeybees has often posed a problem. The consequent use of the advantages offered by honeybees over other test systems ensure a rapidly increasing body of most exciting studies, re-establishing its position as a most rewarding study organism in both physiological and genetical research.
Methods for distinguishing phylogenetic signal from noise are employed to resolve conflicting phylogenies produced by 3 separate morphological and molecular data sets for the 6 species of the honey bee genus Apis. Morphological data from larvae are employed for the 1st time. The results support the phylogeny produced by separate analyses of the morphological and mitochondrial 16s ribosomal DNA (rDNA) data sets, and contradict the result produced by the mitochondrial COII data set. Additionally, these results support previous conclusions that 16s rDNA sequence data are more informative for species-level rather than tribal-level relationships in the corbiculate bees. Resolution of Apis phylogeny permits for the 1st time unambiguous reconstruction of the ancestral states for several important behavioral characters in honey bee evolution.
The history of the systematics of the Asian honeybee species is given, including discussions of paradigm shifts in the definition
of species as earlier taxonomic methods evolved and were gradually replaced by an emphasis on populations, the statistical
distributions of morphological characters and the reconstruction of evolutionary lineages. The limits of Asian species have
been defined using principal component analysis, discriminant analysis, cluster analyses and nearest neighbour procedures,
together with DNA characteristics, behaviour and nesting. This review presents a unified and coherent account of the Asian
honeybees, based on the advances of the last three decades.
This bibliography was compiled from over 3,550 references covering the period 1787 to early 2010. References were obtained
through iterative searches into older sources and continued until a cul-de-sac was reached. Particular effort was expended
in obtaining translations of works in Chinese, Japanese, Thai, Korean, Vietnamese and Russian. Most of the publications cited
here have not been captured by any of the enormous websites or search engines. This bibliography is the most complete of its
kind available. Bibliographic entries are presented with significant keywords relating to the specific biological trait, for
example, swarming, chromosomes, wax and so on as well by scientific names and countries of origin.
This bibliography of the published literature on Apis dorsata and A. laboriosa was compiled from 988 references written by 934 authors published in 290 periodicals, conference proceedings, theses, reports and books covering the period 1793–2005. The literature shows a balance between the applied aspects of beekeeping and basic honeybee biology for A. dorsata; but for A. laboriosa it reflects more basic biology.
A physical map of mitochondrial DNA (mtDNA) of the honeybee (Apis mellifera L) has been established with 17 restriction enzymes (46 sites). Elements of the genic map have been inferred from sequence data. The superimposition of both maps indicate that the gene order is quite similar between honeybee and Drosophila. The total length of the mitochondrial genome falls between 16 500 and 17 000 bp. This range is due to several regions exhibiting length polymorphisms. Two of them overlap with the control region, but a third one is unexpectedly located between the CO-I and CO-II genes. This last polymorphism is explained by the occurrence of variable numbers of 2 related sequences, called P and Q, which arose through tandem duplication. Sequence data from 3 regions of the mtDNA genome can be used to infer a phylogenetic tree for 4 Apis species: the resulting tree topology, (florea(dorsata(cerana,mellifera))), confirms the phylogeny based on morphometry and behavior. The mtDNA variability of Apis mellifera indicates 3 major lineages: African colonies (lineage A) including intermissa, adansonii, scutellata, capensis and monticola subspecies; mellifera colonies (lineage M); ligustica, carnica and caucasica colonies (lineage C). This distribution is very similar to the 3 evolutionary branches inferred from morphometric analysis by Ruttner. The main difference concerns the branch M which, according to Ruttner, includes also intermissa and iberica. From an Asian origin, 3 evolutionary branches colonized respectively northern Europe (M), the north-Mediterranean region (C) and Africa (A). Based on the Drosophila evolutionary rate, this divergence would have occurred between 300 000 and 1300 000 bp.
This bibliography of the literature on Apis florea Fabricius was compiled from 791 references, written by 774 authors and published in 212 different periodicals, conference proceedings, theses, reports, books and patents covering the period 1787–2004. The literature shows greater strength in the applied aspects of beekeeping than in basic biology. Growth of the literature on A. florea has been exponential over the past five decades.
The complete nucleotide sequence of the mitochondrial cytochrome oxidase II (COII) gene was determined for five species of the honeybee (Genus: Apis): A. andreniformis, A. cerana, A. dorsata, A. florea, and A. koschevnikovi; these were then compared to the known sequence of the A. millifera gene from Crozier et al. (1989, Mol. Biol. Evol., 6: 399-411) and the wasp Excristes roborator (Liu and Beckenbach, 1992, Mol. Phylogenet. Evol., 1:41-52). Phylogenetic relationships were derived using the parasimony methods DNAPARS and PROTPARS of Felsenstein ("PHYLIP Manual Version 3.4, "University Herbarium, Univ. of California, Berkeley). The results suggest that A. dorsata is the most ancestral species, followed by the branching of A. florea/A. andreniformis and A. koschevnikovi, and then A. mellifera and A. cerana. This inference differs from the currently accepted view that considers the A. florea/A. andreniformis line to be the most ancestral.
A mitochondrial DNA region encompassing part of the NADH dehydrogenase subunit 2 and isoleucine transfer RNA genes was PCR amplified, cloned, and sequenced for 14 morphometrically identified Apis mellifera subspecies and the New World "Africanized" honeybee. Twenty different haplotypes were detected and phylogenetic analyses supported the existence of 3 or 4 major subspecies groups similar to those based on morphometric measurements. However, some discrepancies are reported concerning the subspecies composition of each group. Based on the sequence divergence of Drosophila (2% per Myr) we found that the four lineages may have diverged around 0.67 Myr. The variability found in this region enables us to infer phylogenetic relationships and test hypotheses concerning subspecies origin, dispersion, and biogeography.
Two different genomic regions (ND2 mitochondrial gene and EF1-alpha intron) were PCR amplified, cloned and sequenced for the ten known honey bee species collected within their natural range distribution. DNA sequences were analyzed using parsimony, distance and maximum likelihood methods to investigate phylogenetic relationships within Apis. The phylogenetic analyses strongly supported the basic topology recoverable from morphometric analysis, grouping the honey bees into three major clusters: giant bees (A. dorsata, A. binghami, and A. laboriosa), dwarf bees (A. andreniformis and A. florea), and cavity-nesting bees (A. mellifera, A. cerana, A. koschevnikovi, A. nuluensis, and A. nigrocincta). However, the clade of Asian cavity-nesting bees included paraphyletic taxa. Exemplars of Apis cerana collected from divergent portions of its range were less related to each other than were sympatric A. cerana, A. nuluensis, and A. nigrocincta taxa. Nucleotide sequence divergence between allopatrically distributed western (A. mellifera) and eastern (A. cerana, A. koschevnikovi, A. nigrocincta, and A. nuluensis) cavity-nesting species, around 18% for the mitochondrial gene and 10-15% for the nuclear intron, suggested an earlier divergence for these groups than previously estimated from morphometric and behavioral studies. This latter finding neccessitates reevaluation of the hypothesized origin of extant European, African, and west Asian Apis mellifera. Sequence divergence between A. laboriosa and A. dorsata was consistent with behavioral data and supports the species status of A. laboriosa.
We characterized Apis mellifera in both native and introduced ranges using 1136 single-nucleotide polymorphisms genotyped in 341 individuals. Our results
indicate that A. mellifera originated in Africa and expanded into Eurasia at least twice, resulting in populations in eastern and western Europe that
are geographically close but genetically distant. A third expansion in the New World has involved the near-replacement of
previously introduced “European” honey bees by descendants of more recently introduced A. m. scutellata (“African” or “killer” bees). Our analyses of spatial transects and temporal series in the New World revealed differential
replacement of alleles derived from eastern versus western Europe, with admixture evident in all individuals.
Starch gel electrophoresis utilizing different types of substrates and inhibitors made it possible to detect several esterases in crude extracts of Apis mellifera. Our results suggest that there are six Apis mellifera esterase isozymes (esterases 1-6) that differ not only in electrophoretic mobility but also in substrate specificity and inhibition properties. Some of the esterase isozymes are controlled by more than one allele. The frequency of these genetic variants was analyzed in four populations of Apis mellifera from several localities. Esterases 1, 2, and 4 do not exhibit developmental changes, but the electrophoretic profile of esterases 3, 4, and 6 varies during ontogenetic development.
For discrete characters whose ancestral states are known, the prescriptions of Hennig are well-defined, but their applicability only when there is no incompatibility between different characters has led to the elaboration of a number of methods for dealing with incompatibilities. One category is the parsimony methods, which choose that phylogeny on which the fewest changes of character state need be assumed. Another is the compatibility methods, which choose that phylogeny which is perfectly compatible with the largest number of characters, irrespective of how many changes need be assumed in other characters. Other approaches include the use of phenetic clustering algorithms and methods fitting trees to similarity or distance matrices. The biological assumptions and statistical behavior of each method are discussed.-from STAR, 20(14), 1982
Six colonies of Apis mellifera mellifera from Norway were electrophoretically examined at twenty-three enzyme loci. Malate dehydrogenase and malic enzyme are polymorphic. Malic enzyme, previously reported to be monomorphic in the honeybee, possesses two allozymes. The lack of malic enzyme variation in the US is probably due to founder effect associated with the introduction of the honeybee into the New World.
BIOSYS-1 is a FORTRAN IV program designed to aid biochemical population geneticists and systematists in the analysis of electrophoretically detectable allelic variation. It can be used to compute allele frequencies and genetic variability measures, to test for deviation of genotype frequencies from Hardy-Weinberg expectations, to calculate F-statistics, to perform heterogeneity chi-square analysis, to calculate a variety of similarity and distance coefficients, and to construct dendrograms using cluster analysis and Wagner procedures. The program, documentation, and test data are available from the authors.
Despite a century of evolutionary theory, it has only been in the last few decades that clearly defined procedures for inferring phylogenies were stated. For discrete characters whose ancestral states are known, the prescriptions of Hennig are well-defined, but their applicability only when there is no incompatibility between different characters has led to the elaboration of a number of methods for dealing with incompatibilities. One category is the parsimony methods, which choose that phylogeny on which the fewest changes of character state need be assumed. Another is the compatibility methods, which choose that phylogeny which is perfectly compatible with the largest number of characters, irrespective of how many changes need be assumed in other characters. Other approaches include the use of phenetic clustering algorithms and methods fitting trees to similarity or distance matrices. Each method has different implicit assumptions concerning the biology of the characters and the information available in the data. Standard statistical approaches such as maximum likelihood can be used to obtain methods whose properties are known, and for which one can determine the amount of uncertainty in the resulting estimates of the phylogeny. It is by viewing the problem as a statistical one that we can place all these methods in a common framework, within which their behavior and assumptions can be compared. It is essential that an attempt be made to understand the biological assumptions and statistical behavior of each method.
A statistical method is developed for estimating the number of gene differences and evolutionary time of a pair of species from electrophoretic data on protein identity. This method is applied to the Drosophila data available. It is shown that the evolutionary time for a pair of nonsibling species in Drosophila is on the average three times longer than that for a pair of sibling species. It is also shown, under certain assumptions, that pairs of recent sibling species differ in about one to two amino acids per protein, and it is estimated that 500,000 years were required to establish such a difference.
it was observed that interspecific copula resulted in severe injury to the ceraqueen. After instrumental insemination of both melli/era and cerana queens with heterospecific semen, normal fertilization and cleavage of the eggs were observed. During the blastula stage, however, development stopped and finally ended in a complete breakdown. These observations are discussed from the point of view of the evolution of the two aforementioned
Motoo Kimura, as founder of the neutral theory, is uniquely placed to write this book. He first proposed the theory in 1968 to explain the unexpectedly high rate of evolutionary change and very large amount of intraspecific variability at the molecular level that had been uncovered by new techniques in molecular biology. The theory - which asserts that the great majority of evolutionary changes at the molecular level are caused not by Darwinian selection but by random drift of selectively neutral mutants - has caused controversy ever since. This book is the first comprehensive treatment of this subject and the author synthesises a wealth of material - ranging from a historical perspective, through recent molecular discoveries, to sophisticated mathematical arguments - all presented in a most lucid manner.
The authors have developed a statistical method for describing relationships among species and higher systematic categories. The method is based on an analysis of a matrix of species-×-species correlation coefficients computed from a large number of characters for each species and has been called the weighted variable group method. Diagrams of relationships resembling phylogenetic trees are constructed and, on the assumption that morphological resemblance is with certain exceptions inversely related to amount of evolutionary divergence, phylogenetic relationships may be hypothesized. The method is applied to a group of four genera of bees in the family Megachilidae. One hundred and twenty-two characters were studied for 97 species of this group. Relationships between the species in this study as visualized by Michener before the statistical analysis are compared with relationships as found in the analysis by Sokal. Good agreement between the two sets of data is found. The few exceptions to this rule are discussed. Some changes in the classification of these bees are made as a result of the analysis and two new subgenera are named. However, the paper serves primarily to illustrate a method that can be used to remove some of the subjective bias from taxonomy.
Thirty-nine colonies of commercial and feral Apis mellifera L. from throughout the United States were surveyed for electrophoretic variation of enzymes known to be polymorphic in the species. These data and those from other studies on U.S. and European honey bees were analyzed to determine the population genetic effects of the bottleneck that accompanied the introduction of this Old World species into the United States. When U.S. populations are compared with those of the Old World, a loss of variation is evident in both the number of polymorphic enzymes and the number of allozymes per polymorphic locus. Evidence is presented for the existence in the United States of a feral gene pool containing an allozyme (Mdh80) common only in A. mellifera mellifera, the first European race to be introduced in the United States. The Mdh100 allozyme also was found in relatively high frequency in feral and commercial colonies and cannot be considered “diagnostic of Africanization in New World honey bees or as a unique character of any African race of A. mellifera, unless a considerable Africanization process has already occurred in the United States.
Thesis (Ph.D.)--University of California, Davis.
Typescript. Thesis (Ph. D.)--University of California, Davis, 1976. Bibliography: leaves 37-40. Microfilm.
An electrophoretic analysis of Rhagoletis (Diptera: Tephritidae) phylogeny. Syst. Zool., 31: 136–155.—Electrophoretic analysis of Rhagoletis (Diptera: Tephritidae) has been performed to gain insight into phylogenetic relationships in the genus, and to determine
whether Rhagoletis species have cospeciated with their host plants. The electrophoretic phylogeny differs from both the conventional classification
and the assumed phylogeny of the host plants. Most of the changes suggested in the conventional classification are compatible
with morphological data. The differences in the phylogenies of Rhagoletis and its host plants are not, however, likely to be resolved by further systematic analysis of the host plants. In addition
to these topics, we also discuss the electrophoretic phylogeny with respect to the biogeography of the genus.
Starch gel electrophoresis of extracts of Apis mellifera indicates that genetic variability exists for the enzyme cytoplasmatic malate dehydrogenase (E.C. 1.1.1.37). Analysis of individuals throughout development indicates that the isozyme patterns are identical for larvae and adults and suggests a dimeric structure for the molecule. The isozyme pattern observed in pupae is more complex than that of larvae and adults and may be due to an additional pupalspecific MDH gene being expressed or to an epigenetic modification of the isozymes. Forty-three colonies with artificially inseminated queens were used to study the Mendelian pattern of inheritance. The data revealed that the MDH isozymes are encoded by three alleles, Mdh-1
A
, Mdh-1
B
, and Mdh-1
C
. The frequency of the Mdh-1 alleles is different in two analyzed subspecies, A. m. adansonii (African bees) and A. m. ligustica (Italian bees), with Mdh-1
A
and Mdh-1
B
in the African bees being 0.768 and 0.202, respectively. For the Italian bees, these frequencies are 0.136 and 0:154, respectively.
Adult workers of the honey bee, Apis mellifera ligustica, from Italy were assayed for enzyme polymorphism using a variety of electrophoretic conditions. Three polymorphic enzyme systems are described, two of which, malic enzyme and an esterase, were previously unknown in indigenous A. m. ligustica. In addition, a new allozyme for the Mdh locus is reported.
Data for Apis mellifera indicate that the P-3 proteins and one esterase enzyme are controlled by two genes, P-3 and Est, with two alleles each. The frequency of the P-3 alleles is different in the two subspecies (adansonii and ligustica), that for P-3F in Italian bees being 46.9% and in African 0.5%. The frequency of EstF is 2.8% in both populations.
The Est locus has two codominant alleles and the locus P-3 has two incompletely dominant alleles; the heterozygote P-3S/P-3F shows only an intermediate band. The two loci are not genetically linked.
ALTHOUGH bees constitute excellent material for genetic and developmental studies, no investigations of biochemical genetics at the protein level have been reported. This article describes two such systems.
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