Population divergence in the ontogenetic trajectories of foliar terpenes of a Eucalyptus species

Background and Aims The development of plant secondary metabolites during early life stages can have significant ecological and evolutionary implications for plant–herbivore interactions. Foliar terpenes influence a broad range of ecological interactions, including plant defence, and their expression may be influenced by ontogenetic and genetic factors. This study investigates the role of these factors in the expression of foliar terpene compounds in Eucalyptus globulus seedlings.Methods Seedlings were sourced from ten families each from three genetically distinct populations, representing relatively high and low chemical resistance to mammalian herbivory. Cotyledon-stage seedlings and consecutive leaf pairs of true leaves were harvested separately across an 8-month period, and analysed for eight monoterpene compounds and six sesquiterpene compounds.Key Results Foliar terpenes showed a series of dynamic changes with ontogenetic trajectories differing between populations and families, as well as between and within the two major terpene classes. Sesquiterpenes changed rapidly through ontogeny and expressed opposing trajectories between compounds, but showed consistency in pattern between populations. Conversely, changed expression in monoterpene trajectories was population- and compound-specific.Conclusions The results suggest that adaptive opportunities exist for changing levels of terpene content through ontogeny, and evolution may exploit the ontogenetic patterns of change in these compounds to create a diverse ontogenetic chemical mosaic with which to defend the plant. It is hypothesized that the observed genetically based patterns in terpene ontogenetic trajectories reflect multiple changes in the regulation of genes throughout different terpene biosynthetic pathways.     

Phylogenetic divergence of fish and mammalian metallothionein: relationships with structural diversification and organismal temperature

Metallothioneins (MTs) are nonenzymatic low molecular weight proteins, that play an important role in the homeostasis and detoxification of heavy metals in a large variety of organisms. These proteins are endowed with striking features, including an unusual amino acid composition characterized by the presence of 20 cysteines out of a total of 60 residues and absence of secondary structure elements. It is generally accepted that MTs underwent few modifications during evolution because of these structural and functional constraints. Such a conclusion is founded on the studies carried out mostly on MTs of mammalian origin. For such a reason, we have decided to compare the MTs of homeothermic and poikilothermic organisms, such as mammals and fish, with the specific aim to put in relation phylogenetic divergence and structural/functional adaptation to temperature. We have included in our analysis also Antarctic Notothenioids, a fish group characterized by genetic isolation and cold-adaptation to a particular harsh environment. We have determined the average hydropathic index of ancestral MT sequences and used them to infer the temperatures of the environment housing the hypothetical ancestor organisms. Finally, we have derived phylogenetic relationships of MT molecules from the pairwise comparison of their three-dimensional structures.     

Shift in the isoelectric-point of milk proteins as a consequence of adaptive divergence between the milks of mammalian species

Background

Milk proteins are required to proceed through a variety of conditions of radically varying pH, which are not identical across mammalian digestive systems. We wished to investigate if the shifts in these requirements have resulted in marked changes in the isoelectric point and charge of milk proteins during evolution.

Results

We investigated nine major milk proteins in 13 mammals. In comparison with a group of orthologous non-milk proteins, we found that 3 proteins κ-casein, lactadherin, and muc1 have undergone the highest change in isoelectric point during evolution. The pattern of non-synonymous substitutions indicate that selection has played a role in the isoelectric point shift, since residues that show significant evidence of positive selection are much more likely to be charged (p = 0.03 for κ-casein; p < 10^-8 for muc1). However, this selection does not appear to be solely due to adaptation to the diversity of mammalian digestive systems, since striking changes are seen among species that resemble each other in terms of their digestion.

Conclusion

The changes in charge are most likely due to changes of other protein functions, rather than an adaptation to the different mammalian digestive systems. These functions may include differences in bioactive peptide releases in the gut between different mammals, which are known to be a major contributing factor in the functional and nutritional value of mammalian milk. This raises the question of whether bovine milk is optimal in terms of particular protein functions, for human nutrition and possibly disease resistance. This article was reviewed by Fyodor Kondrashov, David Liberles (nominated by David Ardell), and Christophe Lefevre (nominated by Mark Ragan).     

Comparison of cranial ontogenetic trajectories among great apes and humans

Molecular data suggest that humans are more closely related to chimpanzees than either is to the gorillas, yet one finds the closest similarity in craniofacial morphology to be among the great apes to the exclusion of humans. To clarify how and when these differences arise in ontogeny, we studied ontogenetic trajectories for Homo sapiens, Pan paniscus, Pan troglodytes, Gorilla gorilla and Pongo pygmaeus. A total of 96 traditional three-dimensional landmarks and semilandmarks on the face and cranial base were collected on 268 adult and sub-adult crania for a geometric morphometric analysis. The ontogenetic trajectories are compared by various techniques, including a new method, relative warps in size-shape space. We find that adult Homo sapiens specimens are clearly separated from the great apes in shape space and size-shape space. Around birth, Homo sapiens infants are already markedly different from the great apes, which overlap at this age but diverge among themselves postnatally. The results suggest that the small genetic differences between Homo and Pan affect early human ontogeny to induce the distinct adult human craniofacial morphology. Pure heterochrony does not sufficiently explain the human craniofacial morphology nor the differences among the African apes.     

Hydrologic regimes as potential drivers of morphologic divergence in fish

Fishes often exhibit phenotypic divergence across gradients of abiotic and biotic selective pressures. In streams, many of the known selective pressures driving phenotypic differentiation are largely influenced by hydrologic regimes. Because flow regimes drive so many attributes of lotic systems, we hypothesized fish exhibit phenotypic divergence among streams with different flow regimes. We used a comparative field study to investigate the morphological divergence of Campostoma anomalom (central stonerollers) among streams characterized by highly variable, intermittent flow regimes and streams characterized by relatively stable, groundwater flow regimes. We also conducted a mesocosm experiment to compare the plastic effects of one component of flow regimes, water velocity, on morphology of fish from different flow regimes. We observed differences in shape between flow regimes likely driven by differences in allometric growth patterns. Although we observed differences in morphology across flow regimes in the field, C. anomalum did not exhibit morphologic plasticity in response to water velocity alone. This study contributes to the understanding of how complex environmental factors drive phenotypic divergence and may provide insight into the evolutionary consequences of disrupting natural hydrologic patterns, which are increasingly threatened by climate change and anthropogenic alterations.     

Genome divergence during evolutionary diversification as revealed in replicate lake-stream stickleback population pairs

Evolutionary diversification is often initiated by adaptive divergence between populations occupying ecologically distinct environments while still exchanging genes. The genetic foundations of this divergence process are largely unknown and are here explored through genome scans in multiple independent lake-stream population pairs of threespine stickleback. We find that across the pairs, overall genomic divergence is associated with the magnitude of divergence in phenotypes known to be under divergent selection. Along this same axis of increasing diversification, genomic divergence becomes increasingly biased towards the centre of chromosomes as opposed to the peripheries. We explain this pattern by within-chromosome variation in the physical extent of hitchhiking, as recombination is greatly reduced in chromosome centres. Correcting for this effect suggests that a great number of genes distributed widely across the genome are involved in the divergence into lake vs. stream habitats. Analyzing additional allopatric population pairs, however, reveals that strong divergence in some genomic regions has been driven by selection unrelated to lake-stream ecology. Our study highlights a major contribution of large-scale variation in recombination rate to generating heterogeneous genomic divergence and indicates that elucidating the genetic basis of adaptive divergence might be more challenging than currently recognized.     

Population genetic divergence as consequence of past range expansion of the smooth hammerhead shark Sphyrna zygaena

Hydrobiologia - We investigated the roles of a benthivorous (Prochilodus brevis, Steindachner 1875) and a planktivorous (Oreochromis niloticus, Linnaeus, 1758) fish in translocating phosphorus from...     

Changes in the lower trophic levels as a consequence of the level of fish manipulation in the ponds

Cyprinid fish of different mature age classes (3+ -4+) and stocks (100, 300 and 500 kg/ha) were introduced into each of three experimental ponds with area of 0.3 ha (average depth ca 1.7 m) while the fourth pond was left free of fish. Bream (Abramis brama L.), white bream (Blicca bjoerkna L.) and roach (Rutilus rutilus L.) made up 75% of the total cyprinid biomass, with wild carp (Cyprinus carpio L.) as the remaining 25%. The introduced fish spawned successfully. The high (above 300 kg/ha) planktivorous and benthivorous fish stocks resulted in several qualitative and quantitative alterations of the food chain structure in our simulation pond experiments. These alterations must primarily be assigned to changes caused by both the zooplanktivory and benthivory nature of the stocked fish populations. At the higher levels of fish biomass, Secchi depth was influenced significantly by chlorophyll-a concentration. Most of the variance in suspended solids concentration could be explained by the biomass ratio of the mature benthivorous fish. There was a clear shift in algal cell size in the ponds with the higher fish stocks: ponds with more fish had larger cells later in the summer. The relative influence of young cyprinid fish on crustaceans species composition and biomass, and mature populations on benthic fauna abundance and biomass, was sufficiently greater at higher (300–500 kg/ha) fish stock rates.     

Household chagasic endemia detected as a consequence of a case of congenital infection]

During a study on prevalence of parasitic and viral serological markers in pregnant adolescents, a 17-year-old primipara from Polpaico, village near Santiago, gave birth to a normal male newborn in a Santiago hospital. As both of them presented positive an indirect hemagglutination test (IHAT) for Chagas' disease and the corresponding xenodiagnosis (XD), the infection in the infant was considered to be acquired through the placental route. According to recent epidemiological surveys Polpaico is an endemo-enzootic chagasic rural settlement, where 14.7% of dwellings were infested with Triatoma infestans, while triatominae, persons and domestic mammals were found infected with Trypanosoma cruzi. One month later the adolescent mother, her son and other 11 consanguineous members of the family were visited in their homes in order to submit each of them to a physical examination and to IHAT for Chagas' disease, and XD to those whose IHAT resulted positive. Thus, in 7 (53.8%) the IHAT was positive and in 4 (57.1%) out of these 7 presented positive the XD, results that as a whole yielded a household chagasic endemics.     

Morphological convergence as a consequence of extreme functional demands: examples from the feeding system of natricine snakes

Despite repeated acquisitions of aquatic or semi-aquatic lifestyles revolving around piscivory, snakes have not evolved suction feeding. Instead, snakes use frontally or laterally directed strikes to capture prey under water. If the aquatic medium constrains strike performance because of its physical properties, we predict morphological and functional convergence in snakes that use similar strike behaviours. Here we use natricine snakes to test for such patterns of convergence in morphology and function. Our data show that frontal strikers have converged on a similar morphology characterized by narrow elongate heads with a reduced projected frontal surface area. Moreover, simple computational fluid dynamics models show that the observed morphological differences are likely biologically relevant as they affect the flow of water around the head. In general, our data suggest that the direction of evolution may be predictable if constraints are strong and evolutionary solutions limited.     

The morphofunctional transformation of skeletal muscle as a consequence of electroneurostimulation training]

By means of histological and histochemical methods slices of biopsies of the canine musculus latissimus dorsi have been investigated after electroneurostimulation for three months through the thoracodorsal nerve in situ and after cutting its initial part. Frequency of contractions increases gradually from 30 up to 80 per 1 min every 2 weeks. The preparations are stained with hematoxylin--eosin. Histochemical reactions for adenosinetriphosphatase (ATPase) (incubational medium pH 4.3 and 10.3) and for acetylcholinesterase (AChE) are performed. The muscle, stimulated in situ, preserves its normal structure. Transformation of muscle contractions from fast to slow, as well as increasing AChE activity in myoneural synapses are revealed. The muscle, stimulated after its cutting at its beginning, is subjected to adipose dystrophy. Activity of ATPase and AChE do not differ from that in the control.     

Patterns of ontogenetic evolution in perianth diversification of Besseya (Scrophulariaceae)

Species of Besseya have perianth diversity centered largely in meristic variation, extreme corolla diminution, and corolla tube loss. Flowers of Besseya differ from those of their closest relative Synthyris in having a bilabiate corolla. The avenues of ontogenetic evolution that were important in creating these aspects of perianth diversity were explored by optimizing developmental transformations on cladograms. Three different evolutionary transformations were important in the diversification of calyx ontogenies in Besseya, including the derived reciprocal substitution of ontogenetic states in B. bullii, labile abaxial lobe expression in B. wyomingensis, and a heterotopic novel substitution in B. oblongifolia. The bilabiate corolla of Besseya arose via fractionation and localization of the zonal growth expressed in the more plesiomorphic Synthyris. Besseya plantaginea, B. ritteriana, and B. oblongifolia form a group characterized by diminished expression of the anterior corolla lobe. Four alternative scenarios equally explain the ontogenetic transformations underlying extreme corolla diminution and corolla tube loss. Corolla tube loss has been considered rare or impossible among angiosperms, but some alternative scenarios in Besseya demonstrate how this may have occurred via fractionation of zonal growth. Corolla tube loss may have arisen only indirectly from the sympetalous ancestors of Besseya following extreme corolla diminution and the reestablishment of a “full-size” corolla which had a novel developmental pattern that did not include the distribution of zonal growth present in ancestors.     

Morphological innovation, diversification and invasion of a new adaptive zone

How ecological opportunity relates to diversification is a central question in evolutionary biology. However, there are few empirical examples of how ecological opportunity and morphological innovation open new adaptive zones, and promote diversification. We analyse data on diet, skull morphology and bite performance, and relate these traits to diversification rates throughout the evolutionary history of an ecologically diverse family of mammals (Chiroptera: Phyllostomidae). We found a significant increase in diversification rate driven by increased speciation at the most recent common ancestor of the predominantly frugivorous subfamily Stenodermatinae. The evolution of diet was associated with skull morphology, and morphology was tightly coupled with biting performance, linking phenotype to new niches through performance. Following the increase in speciation rate, the rate of morphological evolution slowed, while the rate of evolution in diet increased. This pattern suggests that morphology stabilized, and niches within the new adaptive zone of frugivory were filled rapidly, after the evolution of a new cranial phenotype that resulted in a certain level of mechanical efficiency. The tree-wide speciation rate increased non linearly with a more frugivorous diet, and was highest at measures of skull morphology associated with morphological extremes, including the most derived Stenodermatines. These results show that a novel stenodermatine skull phenotype played a central role in the evolution of frugivory and increasing speciation within phyllostomids.     

Morphological structure in a reef fish assemblage

Two key morphological traits, horizontal gape and eye diameter, were measured in a large representative group of coral reef fishes. These morphological traits were used concurrently to assess their utility in exploring abilities of coral reef fishes at an assemblage level. A total of 1,218 specimens from 181 species found on the Great Barrier Reef were examined. Cryptobenthic fishes were included to provide a broader representation of reef fish groups. In the analyses, a clear morphological distinction was found between nocturnal and diurnal fishes. Nocturnal fishes had larger relative horizontal gapes and relative eye diameters by factors of 1.6 and 1.5, respectively. A bivariate plot separated into quadrants was used to assess the implications of morphological variation. The morphological measures reflected distinct ecological traits in each quadrant. Whilst nocturnal fishes had large relative gapes and eye diameters, diurnal predators and detritivores had the same wide gapes, but small relative eye diameters. Highly selective, visual feeders such as the Chaetodontidae and Pseudochromidae had large eyes and small gapes, whilst non-selective feeders with low visual dependence such as the grazing herbivores (Acanthuridae, Siganidae, etc.) had both small eye diameters and gape sizes. The analysis proved to be robust enough to apply to a wide assemblage, but with enough subtlety to distinguish morphological differences within individual families. The methods used in this study may have broad applications to other fish assemblages, both fossil and extant. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Morphological integration and ontogenetic niche shift: a study of crested newt limbs

This study deals with the ontogenetic and evolutionary aspects of integration patterns in the limbs of crested newt species, which, like most amphibians, have a biphasic life history with two morphologically distinct stages (larval vs. juvenile and adult) that occupy different environments (aquatic vs. terrestrial). We analyzed the structure and pattern of correlation between limb skeletal elements at three ontogenetic stages (larval, juvenile, and adult) of four closely related species that differ in their preferences of aquatic habitats (more terrestrial and more aquatic). We found dynamic changes in the pattern of morphological integration between successive ontogenetic stages, as well as changes over the course of crested newt phylogeny. Generally, equivalent ontogenetic stages of different species of crested newts show higher concordance in the correlation pattern than successive ontogenetic stages within species. Among species, two opposing correlation patterns were observed: in more terrestrial species, homologous limb elements are less correlated and within-limb elements are more correlated; in aquatic species, the reverse pattern occurs. These results indicate that the function seems to be the covariance-generating factor, which has shaped the patterns of morphological integration of crested newt limbs.     

The role of ecological opportunity in shaping disparate diversification trajectories in a bicontinental primate radiation

Exceptional species and phenotypic diversity commonly are attributed to ecological opportunity (EO). The conventional EO model predicts that rates of lineage diversification and phenotypic evolution are elevated early in a radiation only to decline later in response to niche availability. Foregut fermentation is hypothesized to be a key innovation that allowed colobine monkeys (subfamily Colobinae), the only primates with this trait, to successfully colonize folivore adaptive zones unavailable to other herbivorous species. Therefore, diversification rates also are expected to be strongly linked with the evolution of traits related to folivory in these monkeys. Using dated molecular phylogenies and a dataset of feeding morphology, I test predictions of the EO model to evaluate the role of EO conferred by foregut fermentation in shaping the African and Asian colobine radiations. Findings from diversification methods coupled with colobine biogeographic history provide compelling evidence that decreasing availability of new adaptive zones during colonization of Asia together with constraints presented by dietary specialization underlie temporal changes in diversification in the Asian but not African clade. Additionally, departures from the EO model likely reflect iterative diversification events in Asia.