Factors influencing progress toward sympatric speciation

Many factors could influence progress towards sympatric speciation. Some of the potentially important ones include competition, mate choice and the degree to which alternative sympatric environments (resources) are discrete. What is not well understood is the relative importance of these different factors, as well as interactions among them. We use an individual-based numerical model to investigate the possibilities. Mate choice was modelled as the degree to which male foraging traits influence female mate choice. Competition was modelled as the degree to which individuals with different phenotypes compete for portions of the resource distribution. Discreteness of the environment was modelled as the degree of bimodality of the underlying resource distribution. We find that strong mate choice was necessary, but not sufficient, to cause sympatric speciation. In addition, sympatric speciation was most likely when the resource distribution was strongly bimodal and when competition among different phenotypes was intermediate. Even under these ideal conditions, however, sympatric speciation occurred only a fraction of the time. Sympatric speciation owing to competition on unimodal resource distributions was also possible, but much less common. In all cases, stochasticity played an important role in determining progress towards sympatric speciation, as evidenced by variation in outcomes among replicate simulations for a given set of parameter values. Overall, we conclude that the nature of competition is much less important for sympatric speciation than is the nature of mate choice and the underlying resource distribution. We argue that an increased understanding of the promoters and inhibitors of sympatric speciation is best achieved through models that simultaneously evaluate multiple potential factors.     

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain. Received: 24 April 1998 / Accepted: 11 May 1999     

The complex genome and adaptive evolution of polyploid Chinese pepper (Zanthoxylum armatum and Zanthoxylum bungeanum)

Zanthoxylum armatum and Zanthoxylum bungeanum, known as ‘Chinese pepper’, are distinguished by their extraordinary complex genomes, phenotypic innovation of adaptive evolution and species-special metabolites. Here, we report reference-grade genomes of Z. armatum and Z. bungeanum. Using high coverage sequence data and comprehensive assembly strategies, we derived 66 pseudochromosomes comprising 33 homologous phased groups of two subgenomes, including autotetraploid Z. armatum. The genomic rearrangements and two whole-genome duplications created large (~4.5 Gb) complex genomes with a high ratio of repetitive sequences (>82%) and high chromosome number (2n = 4x = 132). Further analysis of the high-quality genomes shed lights on the genomic basis of involutional reproduction, allomones biosynthesis and adaptive evolution in Chinese pepper, revealing a high consistent relationship between genomic evolution, environmental factors and phenotypic innovation. Our study provides genomic resources and new insights for investigating diversification and phenotypic innovation in Chinese pepper, with broader implications for the protection of plants under severe environmental changes.     

Form changes in the sea bass, Dicentrarchus labrax (Moronidae: Teleostei), after acclimation to freshwater: an analysis using shape coordinates

Synopsis Morphological changes in the sea bass, Dicentrarchus labrax (Perciformes: Moronidae), were investigated after an experimental acclimation trial to freshwater. The sea bass is an euryhaline species occurring in the Mediterranean and west Atlantic from 30° N to 55° N. Part of the offspring of a pool of breeders was acclimated to freshwater at 9 months of age while maintaining the original stock in marine water. The effect of acclimation to freshwater over the entire form of the fish was studied through geometric morphometrics (shape coordinates). Changes in the form were shown graphically as landmark displacements though age classes 7,12, 15, 19 and 24 months and were discussed in the light of integrated growth. A significantly faster growth in freshwater was detected. Shape coordinates analyzed through multivariate statistics show that significant differences in shape arise after acclimation to freshwater. Moreover, the uniform component of shape change reveals a major effect of stretching or compression perpendicular to the body axis which is common and conservative in both trials. Differences are discussed in terms of selection and ecophenotypism.     

Evolutionary plasticity in prokaryotes: A panglossian view

Enzyme directed genetic mechanisms causing random DNA sequence alterations are ubiquitous in both eukaryotes and prokaryotes. A number of molecular geneticist have invoked adaptation through natural selection to account for this fact, however, alternative explanations have also flourished. The population geneticist G.C. Williams has dismissed the possibility of selection for mutator activity on a priori grounds. In this paper, I attempt a refutation of Williams' argument. In addition, I discuss some conceptual problems related to recent claims made by microbiologists on the adaptiveness of molecular variety generators in the evolution of prokaryotes. A distinction is proposed between selection for mutations caused by a mutator activity and selection for the mutator activity proper. The latter requires a concept of fitness different from the one commonly used in microbiology.     

The role of below-ground competition during early stages of secondary succession: the case of 3-year-old Scots pine (Pinus sylvestris L.) seedlings in an abandoned grassland

In abandoned or extensively managed grasslands, the mechanisms involved in pioneer tree species success are not fully explained. Resource competition among plants and microclimate modifications have been emphasised as possible mechanisms to explain variation of survivorship and growth. In this study, we evaluated a number of mechanisms that may lead to successful survival and growth of seedlings of a pioneer tree species (Pinus sylvestris) in a grass-dominated grassland. Three-year-old Scots pines were planted in an extensively managed grassland of the French Massif Central and for 2 years were either maintained in bare soil or subjected to aerial and below-ground interactions induced by grass vegetation. Soil temperatures were slightly higher in bare soil than under the grass vegetation, but not to an extent explaining pine growth differences. The tall grass canopy reduced light transmission by 77% at ground level and by 20% in the upper part of Scots pine seedlings. Grass vegetation presence also significantly decreased soil volumetric water content (Hv) and soil nitrate in spring and in summer. In these conditions, the average tree height was reduced by 5% compared to trees grown in bare soil, and plant biomass was reduced by 85%. Scots pine intrinsic water-use efficiency (A/g), measured by leaf gas-exchange, increased when Hv decreased owing to a rapid decline of stomatal conductance (g). This result was also confirmed by δ ¹³C analyses of needles. A summer ¹⁵N labelling of seedlings and grass vegetation confirmed the higher NO₃ capture capacity of grass vegetation in comparison with Scots pine seedlings. Our results provide evidence that the seedlings' success was linked to tolerance of below-ground resource depletion (particularly water) induced by grass vegetation based on morphological and physiological plasticity as well as to resource conservation.     

Molecular mechanism of ERK dephosphorylation by striatal‐enriched protein tyrosine phosphatase

Striatal‐enriched tyrosine phosphatase (STEP) is an important regulator of neuronal synaptic plasticity, and its abnormal level or activity contributes to cognitive disorders. One crucial downstream effector and direct substrate of STEP is extracellular signal‐regulated protein kinase (ERK), which has important functions in spine stabilisation and action potential transmission. The inhibition of STEP activity toward phospho‐ERK has the potential to treat neuronal diseases, but the detailed mechanism underlying the dephosphorylation of phospho‐ERK by STEP is not known. Therefore, we examined STEP activity toward para‐nitrophenyl phosphate, phospho‐tyrosine‐containing peptides, and the full‐length phospho‐ERK protein using STEP mutants with different structural features. STEP was found to be a highly efficient ERK tyrosine phosphatase that required both its N‐terminal regulatory region and key residues in its active site. Specifically, both kinase interaction motif (KIM) and kinase‐specific sequence of STEP were required for ERK interaction. In addition to the N‐terminal kinase‐specific sequence region, S245, hydrophobic residues L249/L251, and basic residues R242/R243 located in the KIM region were important in controlling STEP activity toward phospho‐ERK. Further kinetic experiments revealed subtle structural differences between STEP and HePTP that affected the interactions of their KIMs with ERK. Moreover, STEP recognised specific positions of a phospho‐ERK peptide sequence through its active site, and the contact of STEP F311 with phospho‐ERK V205 and T207 were crucial interactions. Taken together, our results not only provide the information for interactions between ERK and STEP, but will also help in the development of specific strategies to target STEP‐ERK recognition, which could serve as a potential therapy for neurological disorders.

     

Nicotine induces dendritic spine remodeling in cultured hippocampal neurons

Cholinergic neurons in the CNS are involved in synaptic plasticity and cognition. Both muscarinic and nicotinic acetylcholine receptors (nAChRs) influence plasticity and cognitive function. The mechanism underlying nAChR‐induced plasticity, however, has remained elusive. Here, we demonstrate morphological changes in dendritic spines following activation of α4β2* nAChRs, which are expressed on glutamatergic pre‐synaptic termini of cultured hippocampal neurons. Exposure of the neurons to nicotine resulted in a lateral enlargement of spine heads. This was abolished by dihydro‐β‐erythroidine, an antagonist of α4β2* nAChRs, but not by α‐bungarotoxin, an antagonist of α7 nAChRs. Tetanus toxin or a mixture of 2‐amino‐5‐phosphonovaleric acid and 6‐cyano‐7‐nitroquinoxaline‐2,3‐dione, antagonists of NMDA‐ and AMPA‐type glutamate receptors, blocked the nicotine‐induced spine remodeling. In addition, nicotine exerted full spine‐enlarging response in the post‐synaptic neuron whose β2 nAChR expression was knocked down. Finally, pre‐treatment with nicotine enhanced the Ca²⁺‐response of the neurons to glutamate. These data suggest that nicotine influences the activity of glutamatergic neurotransmission through the activation of pre‐synaptic α4β2 nAChRs, resulting in the modulation of spinal architecture and responsiveness. The present findings may represent one of the cellular mechanisms underlying cholinergic tuning of brain function.

     

Genetic and developmental basis for parallel evolution and its significance for hominoid evolution

Greater understanding of ape comparative anatomy and evolutionary history has brought a general appreciation that the hominoid radiation is characterized by substantial homoplasy.^1–4 However, little consensus has been reached regarding which features result from repeated evolution. This has important implications for reconstructing ancestral states throughout hominoid evolution, including the nature of the Pan‐Homo last common ancestor (LCA). Advances from evolutionary developmental biology (evo‐devo) have expanded the diversity of model organisms available for uncovering the morphogenetic mechanisms underlying instances of repeated phenotypic change. Of particular relevance to hominoids are data from adaptive radiations of birds, fish, and even flies demonstrating that parallel phenotypic changes often use similar genetic and developmental mechanisms. The frequent reuse of a limited set of genes and pathways underlying phenotypic homoplasy suggests that the conserved nature of the genetic and developmental architecture of animals can influence evolutionary outcomes. Such biases are particularly likely to be shared by closely related taxa that reside in similar ecological niches and face common selective pressures. Consideration of these developmental and ecological factors provides a strong theoretical justification for the substantial homoplasy observed in the evolution of complex characters and the remarkable parallel similarities that can occur in closely related taxa. Thus, as in other branches of the hominoid radiation, repeated phenotypic evolution within African apes is also a distinct possibility. If so, the availability of complete genomes for each of the hominoid genera makes them another model to explore the genetic basis of repeated evolution.