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Sexual selection for size and symmetry in a diversifying secondary sexual character in Drosophila bipectinata Duda (Diptera: Drosophilidae).

Results of intrapopulation studies of sexual selection and genetic variation and covariation underlying elements of the sex comb of Drosophila bipectinata are presented. The magnitude of the sex comb, a sexual ornament, varies significantly among Australasian populations, motivating research into the evolutionary mechanisms responsible for its incipient diversification. The comb is composed of stout black teeth on the front legs of males arranged in three distinct segments: C1, C2, and C3. Significant sexual selection in field populations in northeastern Queensland, Australia, was detected for increasing C2 and body size, and simultaneously for reducing comb positional fluctuating asymmetry. In contrast, sexual selection was not detected for other comb segments, nor for sternopleural bristle number or symmetry. Selection intensities for C2 and comb positional fluctuating asymmetry were similar in magnitude, and although they were opposite in sign, values across twelve sampling dates, or selection episodes, were uncorrelated. Heritability estimates for C2 were high and significant across years, whereas heritability estimates for comb positional asymmetry were small, and generally nonsignificant. The major sex comb segments (C1 and C2) were significantly and positively correlated genetically, indicating the potential for correlated evolution of these components of the comb under sexual selection. The original finding of a significant positive genetic correlation between the magnitude of this sex trait and its positional asymmetry indicates that the counteracting and independent selection pressures detected could contribute to the maintenance of genetic variation sustaining sexual selection. The study documents the simultaneous presence of sexual selection in nature and of heritable genetic variation underlying expression of the sex comb, fundamental conditions necessary for its adaptive diversification. Drosophila bipectinata may be a valuable model for studies of adaptive diversification and incipient speciation by sexual selection.

Adaptation, Biological↗

Larval habits, host-plant associations, and speciation in nematine sawflies (Hymenoptera: Tenthredinidae).

Adaptive radiations consist of two intertwined processes, diversification of species and diversification of their ecological niches, but it is unclear whether there is a causal link between the processes. In phytophagous insects, ecological diversification mainly involves shifts in host-plant associations and in larval feeding habits (internal or external) on different plant parts, and several observations indicate that speciation is facilitated by host shifts. Data on host use in individual species suggest that internal feeders are less likely to colonize new hosts than external-feeding taxa and, consequently, increases in collective host ranges and species numbers should be slowed down in endophagous lineages. We tested these related hypotheses by using phylogenetic information to reconstruct the evolutionary history of larval resource use in the sawfly subfamily Nematinae, a group of 1000 plus species with a broad range of niches: the subfamily's combined host range includes over 20 plant families, and larvae may feed externally on leaves or needles, or internally, for example, in buds, fruits, leaves, or galls. The results show that: (1) Most internally feeding groups have evolved independently from external-feeding ancestors, but several distinct internal habits have appeared convergently multiple times; (2) Shifts among host taxa are clearly more common than changes in larval habits; (3) The majority of host switches have occurred among phylogenetically close plant groups, but many shifts are manifest among distantly related, ecologically proximate hosts; (4) Although external feeding characteristic of the common ancestor of Nematinae is associated with relatively high rates of host-shifting, internal feeders are very conservative in their host use; (5) In contrast, the effect of endophagy on speciation probabilities is more variable: net speciation rates are lowered in most internal-feeding groups, but a striking exception is found in species that induce galls on Salicaceae. The loose connection between collective host ranges and species diversity provides empirical support for theoretical models suggesting that speciation rates are a function of a complex interplay between "intrinsic" niche width and resource heterogeneity.

Animals↗

Ecological speciation in the East Maui-endemic Dubautia (Asteraceae) species.

Adaptive diversification can be viewed as a balance between the conservative force of interpopulation gene flow and selection for differential environments. In this paper, we examine ecological, morphological, and genetic differentiation in a small clade consisting of four East Maui-endemic species of Dubautia: D. menziesii, D. platyphylla, D. reticulata, and D. waianapanapaensis, in the Hawaiian silversword alliance (Asteraceae). The East Maui clade is apparently recently derived (less than 1 million years ago) and is geographically restricted yet displays significant ecological and morphological differences. We used geographic data from historical herbarium specimens, measurements of plant architecture and leaf morphometrics, and measures of genetic differentiation in both microsatellite and nuclear coding loci to examine the correlation of different forms of divergence in this small species flock. We found overlap in large-scale geographic distributions, significant differentiation in most habitat factors, significant microsatellite differentiation, and many shared alleles at nuclear coding loci suggesting on-going lineage sorting. Despite the presence of apparent hybrids in some populations, microsatellite variation is consistent with isolation among species. Using Mantel tests, we compared the direction and extent of diversification among different datasets, to determine whether ecological/morphological divergence was correlated with genetic divergence. Correlations among different datasets showed that habitat was strongly correlated with plant architecture but not leaf morphology. Taken together, these results indicate that ecological and morphological diversification has driven genetic divergence at rapidly evolving microsatellite loci, whereas there is continuing lineage sorting at neutral sites in nuclear coding loci.

Adaptation, Physiological↗

Micro- and macroevolutionary decoupling of cichlid jaws: a test of Liem's key innovation hypothesis.

The extent to which elements of functional systems can change independently (modularity) likely influences the diversification of lineages. Major innovations in organismal design, like the pharyngeal jaw in cichlid fishes, may be key to a group's success when they relax constraints on diversification by increasing phenotypic modularity. In cichlid fishes, pharyngeal jaw modifications that enhanced the ability to breakdown prey may have freed their oral jaws from serving their ancestral dual role as a site of both prey capture and prey processing. This functional decoupling that allowed the oral jaws to become devoted solely to prey capture has been hypothesized to have permitted the two sets of cichlid jaws to evolve independently. We tested the hypothesis that oral and pharyngeal jaw mechanics are evolutionarily decoupled both within and among Neotropical Heroine cichlids. In the trophically polymorphic species Herichthys minckleyi, molariforms that exhibit enlarged molarlike pharyngeal jaw teeth were found to have approximately 400% greater lower jaw mass compared to H. minckleyi with the alternative papilliform pharyngeal morphology. However, oral jaw gape, lower jaw velocity ratios, anterior jaw linkage mechanics, and jaw protrusion did not differ between the morphotypes. In 40 other Heroine species, there was a weak correlation between oral jaw mechanics and pharyngeal jaw mass when phylogenetic history was ignored. Yet, after expansion of the cytochrome b phylogeny for Heroines, change in oral jaw mechanics was found to be independent of evolutionary change in pharyngeal jaw mass based on independent contrasts. Evolutionary decoupling of oral and pharyngeal jaw mechanics has likely played a critical role in the unparalleled trophic diversification of cichlid fishes.

Animals↗

[Evolutionary aspects of the distribution of Ascaridoidea in mammals (author's transl)].

In an attempt to interpret the distribution of Ascaridoidea in Mammals and Birds, a scheme is proposed, based on the concept of geological periods of evolutionary expansion for parasitic lines. It is assumed that the evolutionary expansion period of Ascaridoidea in Mammals occurred after the diversification of primitive mammals; this would explain the absence of ascarids in these groups. Periods of evolution appear to coincide with the diversification of Carnivora and Sirenia, the first to become invaded; Artiodactyla and Equidae became invaded later, at the time of their diversification. The other cases of parasitism of Mammals by Ascarids are thought to be "captures".

Animals↗

Hospital survival strategies for the 1980s.

Survival strategies for hospitals and hospital pharmacy departments are presented, and one hospital's development into a health-services system is described. Financial and competitive pressures are forcing institutions to develop new strategies for survival. The primary institutional strategy is diversification--both horizontal and vertical. Diversification can assist in the hospital's growth, increase its asset and revenue bases, and lead to the development of a health-care delivery system. Marketing to physicians and developing information systems are also critical strategies under a prospective payment system. Institutions will need to know the cost of providing care for specific diagnosis-related groups. Strategies must be employed to develop incentives to reduce inpatient acute-care services and to increase productivity. Physicians should be involved in all of the institutional strategies. Strategies for the pharmacy department are basically the same. Pharmacy departments should develop programs to reduce drug use (especially antibiotic use) and information systems that show the actual cost of providing services by diagnosis. The major corporate restructuring and diversification efforts of one hospital are described. The future of hospitals and pharmacy departments will depend on the successful application of the strategies discussed.

Diagnosis-Related Groups↗

Preferential VH gene usage in rabbit Ig-secreting heterohybridomas.

B cells from leukemic rabbits preferentially use a single VH gene, VH1, in their VDJ gene rearrangements. To determine whether Ig-secreting B cells from normal rabbits also preferentially use VH1, we generated rabbit X mouse heterohybridomas that stably secreted rabbit Ig that expressed VHa allotypic specificities and analyzed the VH genes used in their VDJ gene rearrangements. We cloned the VDJ genes from nine heterohybridomas, and by comparing the restriction map of the DNA immediately 5' of the translational start site of these VDJ genes to that of the same region 5' of VH1, we showed that eight of the nine heterohybridoma clones use VH1. Comparison of the nucleotide sequences of the eight VH1-using VDJ genes with the nucleotide sequence of germ-line VH1 showed that each of them had somatically diversified. The diversified regions included clustered nucleotide changes and codon insertions and deletions, such as would be expected if the diversification process involved somatic gene conversion. We searched our database of germ-line VH genes for genes that could serve as donors for the gene conversion events, and we identified potential VH donor genes for five regions of diversification. One of these regions of diversification spanned at least 132 bp and included a codon insertion as well as 15 nucleotide changes. The data confirm that Ab diversity may be generated by a somatic gene conversion-like mechanism. The results directly demonstrate that Ig-secreting B cells from normal rabbits preferentially use VH1 in their VDJ gene rearrangements.

Amino Acid Sequence↗

RAG1 and RAG2 in developing rabbit appendix subpopulations.

The appendix of young rabbits is a site of primary heavy chain variable region-gene diversification and B cell selection. Appendix cells from 6- to 9-wk-old rabbits were stained and sorted for surface CD43 and IgM. We found that the CD43+IgM- and double-negative CD43-IgM- cells contained RAG1 transcripts and RAG2 protein. The presence of RAG gene products in appendix raised the possibility that pro-/pre-B cells were present in young rabbit appendix. Although an early suggestion that RAG2 plays a role in variable region-gene diversification by gene conversion in chicken bursa was not supported by studies of RAG2 protein in this tissue, we produced anti-rabbit RAG2 Abs to determine whether RAG2 protein was present in rabbit appendix, where cells that recently underwent gene conversion are found. We detected RAG2 protein in the four subpopulations of rabbit appendix lymphocytes, distinguished by surface CD43 and IgM markers. The appearance of RAG gene products during different stages of B cell maturation may reflect the function of the young rabbit appendix as a site of both B cell development and diversification.

Amino Acid Sequence↗

Breast cancer metastasis-associated genes: role in tumour progression to the metastatic state.

Breast cancer patients usually do not die of their primary cancers; they die of metastatic disease. Thus understanding the progression of breast cancer to the metastatic state and the changes that take place in highly malignant breast cells are important goals that could eventually result in new therapeutic approaches to highly progressive breast disease. Changes in the expression of certain genes or alterations in gene structures and encoded products can result in benign tumour cells progressing to the metastatic state. Experimentally, this has been performed by transferring dominantly acting oncogenes into susceptible cells and then testing the malignant properties of these cells in suitable animal models, but such rapid qualitative changes occur in vivo only rarely, and the natural progression of mammary cells to the metastatic state is thought to occur through a slow stepwise process that can take several years. Some of the slow stepwise changes in mammary cancer progression can be reversible and need not involve dominantly acting oncogenes or tumour suppressor genes, consistent with clinical observations. An important element of the natural progression of mammary tumours to malignancy may be their ability to circumvent microenvironmental controls that regulate growth and cellular diversity, a process that appears to involve mainly quantitative changes in gene expression, resulting in loss of normal cellular regulation. One of the important mechanisms of cellular regulation in epithelial tissues, such as those found in the breast, is mediated by intercellular junctional communication. Alterations in gene expression can result in loss of gap-junctional communication, concomitant with cellular diversification and progression. It is thought that the highly malignant cancer cells that have slowly evolved in vivo with only a few qualitative changes in gene structure have undergone extensive cycles of diversification and the accumulation of several quantitative changes in the expression of various genes that encode products related to malignancy. We have identified some of the genes that are related to progression and metastasis in breast cancer. For example, one of these genes, a novel gene called mta1 (in rodents) or MTA1 (in humans) appears to be involved in mammary cell motility and growth regulation. Thus highly malignant cellular phenotypes can arise rapidly due to specific qualitative changes in critical controlling genes, or more slowly via less critical qualitative genetic changes coupled with other cellular changes, such as loss of intercellular communication, and changes in gene expression, such as in the MTA1 gene, resulting in cellular diversification and ultimately tumour progression to the metastatic state.

Breast Neoplasms↗

Metastasis-Associated genes and metastatic tumor progression.

The morbidity in most cancer patients is not due to their primary cancers; it is due to metastatic disease. Thus understanding the progression of tumors to the metastatic state and the changes that take place in highly malignant cells are important in the development of new therapeutic approaches to diagnose, prognostically assess and treat highly progressive malignancies. Changes in the expression of certain genes or alterations of gene structures and encoded products can result in benign tumor cells progressing to the invasive and metastatic states. This has been shown in the laboratory by transferring dominantly acting oncogenes into susceptible cells and then testing the malignant properties of these cells in vivo. Usually such rapid qualitative changes in malignant state occur only rarely; the natural progression of tumor cells to the invasive or metastatic state occurs through a slow stepwise process of change. Tumor progression, in some instances, can be reversible, involving changes in dominantly acting oncogenes or tumor suppressor genes. The natural progression of tumors to highly malignant states also involves their ability to circumvent host microenvironmental controls that regulate cellular growth and diversity. Quantitative changes in gene expression rather than qualitative changes in gene structure are important in microenvironmental effects on progression. One of the important mechanisms of cellular regulation in epithelial tissues, such as breast epithelium, appears to be mediated by intercellular junctional communication. Changes in gene expression can result in loss of junctional communication, followed by cellular diversification and progression. Highly malignant tumor cells that have slowly evolved in vivo with only a few qualitative changes in gene structure have probably undergone extensive cycles of diversification and have multiple quantitative differences in gene expression. Some of these genes are related to metastasis. For example, we have identified a novel gene called mta1 (rat) or MTA1 (human) that appears to be involved in mammary cell motility and growth regulation. This may be an example of a gene that regulates highly malignant cellular phenotypes. When coupled with other cellular changes, such as loss of intercellular communication, specific changes in gene expression may result in cellular diversification and tumor progression.

Animals↗

Structural and tissue-specific organisation of endocrine Fgf19 and Fgf21 signalling in rainbow trout.

Endocrine fibroblast growth factors (FGF19 subfamily) play a key role in regulating metabolic homeostasis in vertebrates. However, their functional diversification in salmonids remains poorly understood. In this study, we conducted an integrative characterisation of Fgf19 and Fgf21 signalling in rainbow trout (Oncorhynchus mykiss) by combining phylogenetic, structural and expression analyses. Phylogenetic analyses revealed the conservation of single fgf19 and fgf21 genes, despite the extensive expansion of receptors post-Ss4R (salmonid-specific fourth-round whole genome duplication). Structural modelling and molecular dynamics simulations demonstrated the stable interactions of both ligands to multiple Fgfr isoforms, with receptor-specific energetic profiles and conserved core interaction residues. Tissue expression profiling revealed clear differences from mammalian models, such as predominant hepatic fgf19 expression and the absence of hepatic fgf21 under basal conditions. In addition, there were complex and tissue-dependent distributions of fgfr and klotho transcripts. These findings support a receptor-driven diversification model of endocrine Fgf signalling in salmonids, suggesting enhanced endocrine plasticity associated with the retention of receptors following post-genomic duplication. Taken together, our findings provide new insights into the structural and regulatory organisation of endocrine Fgf signalling, as well as its potential role in metabolic regulation in rainbow trout.

Animals↗

The rostro-caudal position of cardiac myocytes affect their fate.

During chick embryogenesis, cells destined to form cardiac myocytes are located within the primitive streak at stage 3 in the same relative anterior-posterior distribution as in the prelooped heart. The most rostral cells contribute to the extreme anterior pole of the heart, the bulbus cordis, and the most caudal to the extreme posterior end, the sinoatrial region. After gastrulation, these cells commit to the myocyte lineage and, retaining their relative positions, migrate to the anterior lateral plate. From stages 5 to 10 they diversify into atrial and ventricular myocytes, with the former located posteriorly and the latter, anteriorly. To determine the effect of a change in the rostro-caudal position of these cells on their diversification, anterior lateral plate mesoderm and the underlying endoderm were cut and rotated 180 degrees along the longitudinal axis, at stages 4-8. The subsequent diversification of these precursor cells into atrial and ventricular myocytes was examined using lineage-specific markers. Our results showed that altering location along the longitudinal axis through stage 6 changed the normal fate of a precursor cell. The orientation of the overlying ectoderm did not alter normal morphogenesis or determination of fate.

Animals↗

Exceptionally long CDR3H region with multiple cysteine residues in functional bovine IgM antibodies.

We analyzed VDJ and VJ rearrangements in IgM-secreting B lymphocytes from a cow infected with bovine leukemia virus (BLV). BLV causes expansion of CD5(+) and IgM(+) B lymphocytes regardless of antigen specificity. The data showed that single point mutations contribute to the diversification of IgM antibodies. The most striking observation, however, is that approximately 9% of theVDJ rearrangement in IgM-secreting B cells encode an exceptionally long third complementarity-determining region of the heavy chain (CDR3H; 56 to 61 amino acids) with multiple cysteine residues. Such an exceptionally long CDR3H is the first ever to be documented for an antibody in a species. These VDJ rearrangements encode functional IgM antibodies as some of these show polyspecific reactivity. The presence of even-numbered cysteine residues in the CDR3H may provide hitherto unknown configurational ability to the antigen combining site via intra-CDR3H disulfide bridging. In addition, the VDJ rearrangements encoding exceptionally long CDR3H paired with either novel V(lambda)1 or V(x)1x genes, earlier noted not to be expressed. Overall, these experiments provide evidence that somatic hypermutations and generation of an exceptionally long CDR3H contribute to the diversification of IgM antibodies in cattle.

Amino Acid Sequence↗

Perspectives for crop protection in sustainable agriculture.

The challenge facing agriculture, particularly in developing countries, is to increase productivity without causing ecological damage. An important aspect of sustainable agriculture is the substitution of chemicals and capital with locally grown biological inputs and knowledge. Production of food grains in India has risen steadily since the 1960s, partly through the introduction of high-yielding varieties of rice and wheat together with appropriate agronomic and plant protection practices. Plant breeding programmes have improved cultivated rice by the transfer, from wild species, of genes conferring resistance to viral diseases. Diversification of resistance is sought, to render the hybrids less vulnerable to sudden outbreaks of diseases and insect pests. For wheat, resistance to rusts involves the additive interaction of several gene-encoded traits. Linkage of resistance genes with morphological markers enables the inheritance of the former to be followed easily. It is important that recent advances in molecular genetics are incorporated into integrated pest management programmes. This will require appropriate social organization. Research centres that develop and exploit genetic sources of resistance and varietal diversification should be established.

Agriculture↗

Structure and function of insect olfactory sensilla.

Olfactory sensilla show a large diversification of sensillum types even in the same species. Thus, double-walled and single-walled sensilla with highly different wall pores are usually found on the same antenna, and these may appear in the form of long slender hairs, pore plates or pit pegs. The selective constraints leading to this diversification are evident only in a few cases, e.g. the demand for extreme sensitivity in moth pheromone communication supported the evolution of long sensilla trichodea with high efficiency of capturing odour molecules. The structural diversity continues with the odorant-binding proteins (OBPs) in the sensillum lymph surrounding the sensory dendrites. These proteins may be subdivided into pheromone-binding proteins and two classes of general odorant-binding proteins according to their primary sequence. Different sensilla of the same morphological type may contain different OBPs of the same or of different subclasses. However, OBPs of different subclasses are not co-localized in the same individual sensory hair. The presence of a given OBP is related more to the functional specificity of the receptor cells than to the morphological type of the sensillum, suggesting a role of OBPs in stimulus recognition.

Animals↗

Resolving the "Yucca queretaroensis problem": Phylogenomic analysis of Yucca reveals the identity of an enigmatic species and the origin of an obligate pollination mutualism.

PREMISE: The genus Yucca is a group of ~50 species of woody monocots endemic to the North American arid regions. Their obligate pollination mutualism with yucca moths is considered a "textbook example" of coevolution and is hypothesized to have promoted rapid diversification. However, testing this hypothesis has been difficult due to uncertainty about the placement of a rogue taxon, Yucca queretaroensis, a rare endemic of the Sierra Gorda region of central Mexico. Past work placed this species in different positions within the Agavoideae, producing starkly different age estimates for Yucca (25 to 4 million years). METHODS: We generated new sequence capture data for 353 nuclear genes and for all coding regions of the plastid genome from wild-collected plants and samples included in previous studies to provide a new phylogeny and new age estimate for Yucca. RESULTS: The data presented here suggest that Y. queretaroensis is closely related to other species of Yucca. A relaxed molecular clock analysis of the plastid genome produced an estimated age for the genus of approximately 6.8 million years. CONCLUSIONS: The results resolve a mystery that has bedeviled evolutionary biologists for decades and provide a surprisingly young estimate for the age of Yucca, suggesting rapid diversification. The past difficulties in identifying the correct placement of Y. queretaroensis appear to be the product of laboratory errors, mistakes in field identification, and frequent hybridization with co-distributed taxa. The "Yucca queretaroensis problem" reaffirms the essential role for traditional botanical tools in phylogenomics.

ASTRAL↗

Discerning evolutionary processes in patterns of tamarin (genus Saguinus) craniofacial variation.

Quantitative genetic theory specifies evolutionary expectations for morphological diversification by genetic drift in a monophyletic clade. If genetic drift is responsible for the evolutionary morphological diversification of a clade, patterns of within- and between-taxon morphological variance/covariance should be proportional. We tested for proportionality of within- and between-species craniofacial morphological variation in 12 species of tamarins (genus Saguinus). We found that within- and between-taxon morphological variations across the entire genus were not proportional, and hence not likely to be due to genetic drift alone. The primary deviation from proportionality is that size and size-related shape in the cranium is more variable relative to other aspects of cranial morphology than expected under genetic drift, suggesting differential size selection between the two major clades, the small-bodied and large-bodied tamarins. Within each of these major clades, most of the interspecific variation is consistent with the pattern expected under genetic drift, although specific contrasts may indicate the involvement of differential selection. Morphological distances among taxa do not correspond very closely to the phylogeny derived from mtDNA. In particular, S. oedipus and S. geoffroyi are very distinct morphologically from the rest of the tamarins, although they are phylogenetically the sister clade to a clade containing S. midas and S. bicolor. Morphological similarity is not a good guide to phylogenetic affinity in the tamarins, especially with regard to deeper nodes in the phylogenetic tree.

Animals↗

Origin of multicellular organisms as an inevitable consequence of dynamical systems.

The origin of multicellular organisms is studied by considering a cell system that satisfies minimal conditions, that is, a system of interacting cells with intracellular biochemical dynamics, and potentiality in reproduction. Three basic features in multicellular organisms-cellular diversification, robust developmental process, and emergence of germ-line cells-are found to be general properties of such a system. Irrespective of the details of the model, such features appear when there are complex oscillatory dynamics of intracellular chemical concentrations. Cells differentiate from totipotent stem cells into other cell types due to instability in the intracellular dynamics with cell-cell interactions, as explained by our isologous diversification theory (Furusawa and Kaneko, 1998a; Kaneko and Yomo, 1997). This developmental process is shown to be stable with respect to perturbations, such as molecular fluctuations and removal of some cells. By further imposing an adequate cell-type-dependent adhesion force, some cells are released, from which the next generation cell colony is formed, and a multicellular organism life-cycle emerges without any finely tuned mechanisms. This recursive production of multicellular units is stabilized if released cells are few in number, implying the separation of germ cell lines. Furthermore, such an organism with a variety of cellular states and robust development is found to maintain a larger growth speed as an ensemble by achieving a cooperative use of resources, compared to simple cells without differentiation. Our results suggest that the emergence of multicellular organisms is not a "difficult problem" in evolution, but rather is a natural consequence of a cell colony that can grow continuously.

Algorithms↗