EVOLUTION OF FLORAL TRAITS IN A HERMAPHRODITIC PLANT: FIELD MEASUREMENTS OF HERITABILITIES AND GENETIC CORRELATIONS

Genetic variances, heritabilities, and genetic correlations of floral traits were measured in the monocarpic perennial Ipomopsis aggregata (Polemoniaceae). A paternal half-sib design was employed to generate seeds in each of four years, and seeds were planted back in the field near the parental site. The progeny were followed for up to eight years to estimate quantitative genetic parameters subject to natural levels of environmental variation over the entire life cycle. Narrow-sense heritabilities of 0.2–0.8 were detected for the morphometric traits of corolla length, corolla width, stigma position, and anther position. The proportion of time spent by the protandrous flowers in the pistillate phase (“proportion pistillate”) also exhibited detectable heritability of near 0.3. In contrast, heritability estimates for nectar reward traits were low and not significantly different from zero, due to high environmental variance between and within flowering years. The estimates of genetic parameters were combined with phenotypic selection gradients to predict evolutionary responses to selection mediated by the hummingbird pollinators. One trait, corolla width, showed the potential for a rapid response to ongoing selection through male function, as it experienced both direct selection, by influencing pollen export, and relatively high heritability. Predicted responses were lower for proportion pistillate and corolla length, even though these traits also experienced direct selection. Stigma position was expected to respond positively to indirect selection of proportion pistillate but negatively to selection of corolla length, with the net effect sensitive to variation in the selection estimates. Anther position also was not directly selected but could respond to indirect selection of genetically correlated traits.     

A TRADE-OFF FOR HOST PLANT UTILIZATION IN THE BLACK BEAN APHID,APHIS FABAE

Many studies on insect herbivores have sought to find trade-offs between utilization of alternate host plants, both to understand the prevalence of specialization and to appreciate the likelihood of sympatric speciation due to disruptive selection. To date, few studies have found trade-offs. Seventy-seven clones of the black bean aphid, Aphis fabae, were collected from field sites in East Anglia, U.K., over an area of about 10,000 km². These clones exhibit a trade-off in fitness between two alternative hosts, broad bean and nasturtium. This pattern is maintained in the F2 generation. The predominance of broad bean in the area, the fact that clones were only sampled from one of these two hosts, and the absence of “master-of-all-trades” genotypes after recombination all point to the importance of antagonistic pleiotropy rather than linkage disequilibrium in maintaining this trade-off. It is concluded that this population presents strong evidence for a fundamental trade-off for host utilization.     

Small-scale association measures in epibenthic communities as a clue for allelochemical interactions

The small-scale associations in a rocky subtidal community in the northwestern Mediterranean were studied by a development of the continuous line transect method. This method allowed the overall measurement of non-randomness in interspecific contacts and the assignment of an association index to each species-pair, whose, significance was tested by Monte Carlo procedures. At the same time, the continuous recording allowed the study of the weakening of the interactions with increasing distances. Our purpose was to uncover evidence for allelochemical mechanisms of space occupation and maintenance. A strong non-randomness was found in the interspecific associations. This was mostly due to the interactions of the poecilosclerid sponge Crambe crambe (Schmidt) with its neighbours, especially its negative associations with other sponge species. The strength of the relationships fell drastically over the first few centimeters from the contact borders of the different species. The results pointed strongly to an allelochemical mechanism. The extracts of this sponge featured high bioactivity in laboratory assays, and field experiments demonstrated that the sponge can inhibit the growth of species in the community studied. Standard sampling techniques would have overlooked the spatial structure present in the data. The study emphasizes the need for both contact data and distance data in order to identify the underlying processes reliably. The line transect method provides both types of information easily and allows testing of models and identification of organisms likely to use chemical defenses in space competition. Its use as a preliminary step in studies of chemical ecology might help to detect presumptive allelochemical processes prior to experimental work on the potentially active species.     

Stochastic determinants of assemblage patterns in coral reef fishes: a quantification by means of two models

Synopsis One perspective emphasizing the importance of stochastic processes in determining coral reef fish assemblages implies that there is little organization in species richness, abundance structure, and spatial distribution. We examine the degree to which this perspective is correct by analyzing distribution of fishes on a collection of patch reefs (Discovery Bay, Jamaica). We ask the question whether these patches accumulate species and individuals in a manner consistent with stochastic expectations. To address this question we use two conceptual models, each permitting a different insight. One assumes that fish are distributed stochastically on patches while the other assumes presence of restrictions on fish distribution due to habitat structure. For each conceptual model we use two types of benchmark: we compare observed patterns to those predicted by theoretical models, and we also compare observed patterns to those obtained from a random reallocation of fish individuals to patches. We found that the conceptual model assuming stochastic processes appeared to provide weaker explanation of patterns than the conceptual model that includes restrictions due to habitat structure. Further, and more importantly, we found that (i) the community is shaped by a mixture of stochastic and non-stochastic mechanisms, and (ii) the stochastic assembly processes decrease in importance for species restricted to fewer microhabitat types and sites. Our study therefore indicates that patches do accumulate individuals and species in a manner consistent with stochastic expectations, however, this applies primarily to the habitat generalists (unrestricted species). By the same token, increased habitat specialization by some species imposes constraints on the stochastic model such that it eventually fails.     

Conformation of 4-thio-l-lyxono-1,4-lactone in solution and in the crystalline state

The conformation in ²H₂O of 4-thio-l-lyxono-1,4-lactone (1) was studied by nuclear magnetic resonance spectroscopy, by means of homonuclear (J_1H,1H) and heteronuclear (J_1H,13C) coupling constants. The couplings were directly measured by a two-dimensional heteronucleus-coupled ω₁ hetero-half-filtered proton-proton correlation (HETLOC) experiment, which does not require ¹³C isotopic enrichment. In solution, the thiolactone ring of 1 adopts preferentially the E₃ conformation, and its hydroxymethyl group populates mainly the gt rotamer. The X-ray diffraction data of a single crystal of 1 indicates that also in the solid state the thiolactone ring adopts an E₃ conformation, with a puckering somewhat larger than that observed for aldono-1,4-lactones and furanose rings. The molecules are linked by hydrogen bonds, which form chains. Particularly, O-5 is fully engaged as donor and acceptor in hydrogen bonding and the rotameric conformation of the hydroxymethyl group of 1 is fixed in the tg form.     

The status and ecology of a Juniperus excelsa subsp. polycarpos woodland in the northern mountains of Oman

Juniperus excelsa subsp. polycarpos (K. Koch) Takhtajan is found in mountain areas from Turkey through to India and as an isolated population on Jebel Akhadar in the northern mountains of Oman. Juniperus is one of the dominant plant species in these mountains and a major landscape feature of several proposed National Nature and Scenic Reserves and of Hayl Juwari, a wooded valley at 2250 m altitude proposed as a Botanical Site of Special Interest. Above 2400 m altitude the Juniperus woodlands generally appear to be regenerating and in good condition, both on exposed slopes and in wadis and sheltered gullies, whereas below 2400 m most stands are in poor condition and exhibit few signs of regeneration. If the apparently poor condition of the lower altitude woodlands is due to any long term change in climatic conditions, both tree status and regeneration would be poorer in relatively more xeric habitats. To test this prediction we have carried out a detailed survey of the status and ecology of a 32 ha area of Hayl Juwari, and analysed differences in tree status and regeneration between wadis (relatively more nesic sites) and non-wadi areas (relatively more xeric). Approximately one third of the trees are dead, and an analysis of the height, condition, regeneration, female cone production, preferred germination sites and spatial distribution of trees indicates the importance of topography, hydrology and microclimate for growth. However, although there are relatively greater numbers of dead and poor-condition trees in the more xeric non-wadi habitat, there is no unequivocal evidence that the present distribution of small, sexually immature trees in both habitats could not form a pattern of larger, sexually mature trees similar to that seen today. We speculate, however, that the climate at this altitude may be marginal for the survival of a J. excelsa subsp. polycarpos woodland and that even small increases in climatic stress could imperil the woodland's present status.     

Natural disturbance and tree species coexistence in an old-growth beech - dwarf bamboo forest,southwestern Japan

Abstract. The structure and composition of a cool-temperate old-growth beech (Fagus crenata) - dwarf bamboo (Sasa spp.) forest, partially affected by landslide disturbance, in the Daisen Forest Reserve of southwestern Japan, were investigated in relation to forest floor and canopy conditions. All stems ≥ 4 cm DBH were mapped on a 4-ha plot and analyses were made of population structure, spatial distribution and spatial association of major tree species. The dominant species, F. crenata, which had the maximum DBH among the species present, had the highest stem density. However, for other species, larger-sized species had lower stem density with few smaller stems or saplings, while smaller-sized species had higher stem density with many smaller stems or saplings. Canopy trees of F. crenata were distributed randomly in the plot, while its stems in the other layers and all other species were distributed patchily. Small patches represent gap-phase regeneration. Larger patches correlate with landslide disturbance, difference in soil age, or the presence/absence of Sasa. Cluster analysis for spatial associations among species and stems in the different layers revealed that the forest community consists of several groups. One main group was formed on sites not covered with Sasa. This group contained a successional subgroup (from Betula grossa to Acer mono and/or F. crenata) initiated by landslide disturbance and a subgroup of tree species that avoid Sasa. Another group was formed on sites with mature soils covered largely with Sasa. This contained associations of canopy trees of F. crenata and smaller-sized tree species such as Acanthopanax sciadophylloides and Acer japonicum. It is found that the community of this old-growth beech forest is largely organized by natural disturbance and heterogeneous conditions of the forest floor (difference in soil age and presence/absence of Sasa). The existence of these different factors and the different responses of species to them largely contribute to the maintenance of tree species diversity in this forest.; Keywords: Cluster analysis; Fagus crenata; Forest dynamics; Gap; Landslide; Spatial pattern.     

Oscillatory reaction-diffusion equations on rings

We study the behavior of traveling waves in - systems on both homogeneous and inhomogeneous rings. The stability regions in parameter space of - waves were previously known [15, 19]; the results are extended here. We show the existence of Hopf bifurcations of traveling waves and the stability of the limit cycles born at the Hopf bifurcation for some parameter ranges. Using a Lindstedt-type perturbation scheme, we formally construct periodic solutions of the - system near a Hopf bifurcation and show that the periodic solutions superimposed on the original traveling wave have the effect of altering its overall frequency and amplitude. We also study the - system on an annulus ofvariable width, which does not possess reflection symmetry about any axis. We formally construct traveling waves on this variable-width annulus by a perturbation scheme, and find that perturbing the width of the annulus alters the amplitude and frequency of traveling waves on the domain by a small (order ²) amount. For typical parameter values, we find that the speed, frequency, and stability are unaffected by the direction of travel of the wave on the annulus, despite the rotationally asymmetric inhomogeneity. This indicates that the - system on a variable-width domain cannot account for directional preferences of traveling waves in biological systems.     

Genetic variation in North Africa and Eurasia: Neolithic demic diffusion vs. paleolithic colonisation

The hypothesis that both genetic and linguistic similarities among Eurasian and North African populations are due to demic diffusion of neolithic farmers is tested against a wide database of allele frequencies. Demic diffusion of farming and languages from the Near East should have determined clines in areas defined by linguistic criteria; the alternative hypothesis of cultural transmission does not predict clines. Spatial autocorrelation analysis shows significant gradients in three of the four linguistic families supposedly affected by neolithic demic diffusion; the Afroasiatic family is the exception. Many such gradients are not observed when populations are jointly analyzed, regardless of linguistic classification. This is incompatible with the hypothesis that major cultural transformations in Eurasia (diffusion of related languages and spread of agriculture) took place without major demographic changes. The model of demic diffusion seems therefore to provide a mechanism explaining coevolution of linguistic and biological traits in much of the Old World. Archaeological, linguistic, and genetic evidence agree in suggesting a multidirectional process of gene flow from the Near East in the neolithic. However, the possibility should be envisaged that some allele frequency patterns can predate the neolithic and depend on the initial spread of Homo sapiens sapiens from Africa into Eurasia. © 1994 Wiley-Liss, Inc.     

Trade-offs in responses to host plants within a population of a generalist herbivore,Christoneura rosaceana

Evolutionary constraints on the ability of herbivores to efficiently use a set of phytochemically similar hosts, while maintaining a high performance on phytochemically different hosts, are central in explaining the predominance of host specialization in phytophagous insects. Such feeding trade-offs could be manifested within insect populations as negative genetic correlations in fitness on different host species. We tested the hypothesis that feeding trade-offs were present within a population of the obliquebanded leafroller,Choristoneura rosaceana (Harris). Components of fitness were measured in families originating from an apple orchard that were fed on four host-plant species in the laboratory. Under the conditions of this experiment, all across-host genetic correlations were strongly positive, suggesting that this population comprised true generalists. With the exception of diapausing propensity, the heritability of the fitness components tended to be lower in caterpillars fed on apple leaves than in insects fed other hosts. This suggests a constraint on the selective response of the fitness components in the orchard environment.