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Trophic cascades across ecosystems.

Predation can be intense, creating strong direct and indirect effects throughout food webs. In addition, ecologists increasingly recognize that fluxes of organisms across ecosystem boundaries can have major consequences for community dynamics. Species with complex life histories often shift habitats during their life cycles and provide potent conduits coupling ecosystems. Thus, local interactions that affect predator abundance in one ecosystem (for example a larval habitat) may have reverberating effects in another (for example an adult habitat). Here we show that fish indirectly facilitate terrestrial plant reproduction through cascading trophic interactions across ecosystem boundaries. Fish reduce larval dragonfly abundances in ponds, leading to fewer adult dragonflies nearby. Adult dragonflies consume insect pollinators and alter their foraging behaviour. As a result, plants near ponds with fish receive more pollinator visits and are less pollen limited than plants near fish-free ponds. Our results confirm that strong species interactions can reverberate across ecosystems, and emphasize the importance of landscape-level processes in driving local species interactions.

Animals↗

Genetic engineering of parthenocarpic plants.

Transgenic tobacco and eggplants expressing the coding region of the iaaM gene from Pseudomonas syringae pv. savastanoi, under the control of the regulatory sequences of the ovule-specific DefH9 gene from Antirrhinum majus, showed parthenocarpic fruit development. Expression of the DefH9-iaaM chimeric transgene occurs during flower development in both tobacco and eggplant. Seedless fruits were produced by emasculated flowers. When pollinated, the parthenocarpic plants produced fruits containing seeds. In eggplant, the genetic manipulation allowed fruit set and growth under environmental conditions prohibitive for fruit setting in the untransformed line, which did not set fruit at all. Under normal environmental conditions, production of marketable fruits took place from pollinated and unpollinated transgenic flowers, while flowers of untransformed control plants did produce fruits of marketable size only from fertilized flowers.

Chimera↗

Temporal aspects of the fine-scale genetic structure in a population of Cinnamomum insularimontanum (Lauraceae).

Cinnamomum insularimontanum Hayata (Lauraceae) is an insect-pollinated, broad-leaved evergreen tree with bird-dispersed seeds. We used allozyme loci, Wright's fixation index, spatial autocorrelation statistics (Moran's I), and coancestry measures to examine changes in genetic structure among four age-classes within a recently founded study population (60 x 100 m area) in southern Korea. There were no significant differences in expected heterozygosity among age classes. However, significant genetic differentiation among age classes was detected (P<0.0001). Fixation indices within age classes showed significant deficits of observed heterozygosity, which may be caused by partial selfing. The homogeneity of genetic structure among four age-classes may reflect similar spatial patterns of seed immigration from surrounding populations occurring year after year. Finally, the average Moran's I and coancestry estimates indicated essentially random spatial distributions of alleles for each of the four age-classes and between seedlings and 2-4 year juveniles vs adult trees. These findings are very similar to those observed in the same study area for another member of the Lauraceae, Neolitsea sericea, which has a very similar life history and ecological characteristics (ie, bird-dispersed fruits, insect pollination, and a similar age structure). Together, these results suggest that the fleshy drupes of lauraceous species represent an adaptation to aid in the independent dispersal of seed by birds, which in turn may increase the genetic diversity of founders colonizing new habitats.

Biological Evolution↗

The genetic structure of a columnar cactus with a disjunct distribution: Stenocereus gummosus in the Sonoran desert.

Stenocereus gummosus is a columnar cactus endemic to the Sonoran desert that exhibits a disjunct distribution: it is widely distributed in Baja California and restricted to a small coastal area in mainland Sonora. In this paper, we examine the genetic structure and the mating system of this species in order to explore the origin of the disjunction and describe aspects of the pollination biology. Flowers are nocturnal, pollinated mainly by sphingids and self-incompatible. Polymorphism for allozymes (11 loci) was relatively high (P=75%) but moderate levels of heterozygosity were detected (H(o)=0.103 and H(e)=0.261). Sonoran populations exhibited higher levels of genetic variation than peninsular populations. H(e) declined with latitude when just peninsular and two island populations are included. Substantial levels of inbreeding within populations (f=0.60), moderate differentiation among populations (theta=0.10), and no evidence of isolation by distance were detected. The neighbor-joining phenogram showed Sonoran and island populations nested within peninsular populations. Mainland populations showed greater genetic similarity to island populations, supporting a dispersal hypothesis for the origin of the disjunction. Future studies using DNA markers are suggested in order to better understand the forces that have shaped the genetic structure of this species.

Cactaceae↗

Effects of seed bank disturbance on the fine-scale genetic structure of populations of the rare shrub Grevillea macleayana.

Dispersal in most plants is mediated by the movement of seeds and pollen, which move genes across the landscape differently. Grevillea macleayana is a rare, fire-dependent Australian shrub with large seeds lacking adaptations for dispersal; yet it produces inflorescences adapted to pollination by highly mobile vertebrates (eg birds). Interpreting fine-scale genetic structure in the light of these two processes is confounded by the recent imposition of anthropogenic disturbances with potentially contrasting genetic consequences: (1) the unusual foraging behaviour of exotic honeybees and 2. widespread disturbance of the soil-stored seedbank by road building and quarrying. To test for evidence of fine-scale genetic structure within G. macleayana populations and to test the prediction that such structure might be masked by disturbance of the seed bank, we sampled two sites in undisturbed habitat and compared their genetic structure with two sites that had been strongly affected by road building using a test for spatial autocorrelation of genotypes. High selfing levels inferred from genotypes at all four sites implies that pollen dispersal is limited. Consistent with this, we observed substantial spatial clustering of genes at 10 m or less in the two undisturbed populations and argue that this reflects the predicted effects of both high selfing levels and limited seed dispersal. In contrast, at the two sites disturbed by road building, spatial autocorrelation was weak. This suggests there has been mixing of the seed bank, counteracting the naturally low dispersal and elevated selfing due to honeybees. Pollination between near neighbours with reduced relatedness potentially has fitness consequences for G. macleayana in disturbed sites.

Australia↗

Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: II. Reproductive strategies.

This study compares sexually and asexually produced fruit set, seed production, biomass, germination, and seedling size in Carpobrotus acinaciformis and C. edulis following controlled pollination experiments in order to evaluate the potential role of reproductive traits with respect to the invasive potential of these taxa. C. edulis is slightly agamospermic, completely self-fertile, slightly preferentially self-compatible, experiences no inbreeding depression, and has low hybrid vigour. In contrast, C. acinaciformis does not have reliable agamospermy, is only slightly self-fertile and self-compatible, experiences a slight inbreeding depression, and has a strong hybrid vigour. Both taxa have relatively low, although significantly different germination frequencies, and insignificantly different seedling sizes. Owing to the high performance in hybridisation as compared to all other controlled pollinations in C. acinaciformis, as well as a large amount of previously demonstrated introgression, we refer to the population studied on the island of Bagaud (France) as C. affine acinaciformis. We conclude that both C. edulis and C. affine acinaciformis should be considered as harmful invasive plants in the Mediterranean Basin, the former because of the flexibility of its mating system and high seed production, and the latter because of its strong clonality, high hybrid vigour, and potential for continued introgression from C. edulis genes. These differences require different control strategies, while the avoidance of sympatry is a distinct priority.

Aizoaceae↗

Identification and mode of action of self-compatibility loci in Lolium perenne L.

The two-locus gametophytic incompatibility system in perennial ryegrass (Lolium perenne L.) is not always fully effective: obligate selfing of plants sieves self-compatible pollen mutants, and self-fertility becomes fixed in subsequent generations. Self-compatibility (SC) was investigated in an F2 family. In vitro self-pollinations were analysed and recorded and plants were classified as being either partially or fully compatible. Distorted segregation ratios of markers on linkage group (LG) 5 were found, which indicate the possible presence of a gametophytic SC locus. Interval linkage analysis of pollen compatibility after selfing confirmed that this distortion was due to a locus (T) analogous to the S5 locus of rye. However, even though markers in this region were, on average, less than 1 cM apart, the minimum number of plants possessing the unfavoured allele was never less than 6% for any marker locus. We proved that this was because of the presence of another SC locus, exhibiting gametophytic selection, segregating in this population and identified by interval mapping analysis of compatibility classes of in vitro self-pollinations. This locus was located on LG1, and probably corresponds to the S locus. We show that the T locus, a relic of a multilocus system, functions through interaction with the S locus: F2 segregation of incompatibility phenotypes and linked markers demonstrated that the S/t pollen genotype combination, expected to be compatible on selfing, was sometimes incompatible. Further evidence is presented to show that this interaction must be dependent on yet another locus located on LG2. A prime candidate would be the Z incompatibility locus.

Chromosome Mapping↗

Spatial genetic structure of two sympatric neotropical palms with contrasting life histories.

The spatial genetic structure within sympatric populations of two neotropical dioecious palm species with contrasting life histories was characterized to evaluate the influence of life history traits on the extent of genetic isolation by distance. Chamaedorea tepejilote is a common wind-pollinated arboreal understory palm. Chamaedorea elatior is an uncommon climbing subcanopy palm with entomophilous pollination syndrome. A total of 59 allozyme alleles for C. tepejilote and 53 alleles for C. elatior was analyzed using both unweighted (Iu) and weighted (Iw) Moran's I spatial autocorrelation statistics. The spatial genetic structure detected within these populations is consistent with those reported for highly dispersed plants. A significance test for differences between mean Moran's I-coefficients revealed less spatial genetic structure within the C. tepejilote population than that in the C. elatior population. Adjacent individuals of C. elatior exhibited significant spatial genetic autocorrelation (Iu=0.039, Iw=0.034), indicating a Wright's neighborhood size of about 100 individuals. For C. tepejilote, nonrandom genetic distribution among nearest neighbors was detected, even from small spatial autocorrelation values (Iu=0.008, Iw=0.009), consistent with a neighborhood size of about 300 individuals. For both species, seed dispersal, mortality among life cycle stages, overlapping generations, and contrasting traits of mating and reproduction influence the standing spatial genetic structure within populations.

Arecaceae↗

Effect of variation in herkogamy on outcrossing within a population of Gilia achilleifolia.

The diverse features of floral morphology are often thought to be well-designed mechanisms to manipulate plant mating systems. We evaluated the effectiveness of one such mechanism, anther-stigma separation (herkogamy), in controlling variation in the level of outcrossing among plants in a population of Gilia achilleifolia. Variation in outcrossing rates within populations has the potential to influence the coevolution between inbreeding depression and mating system. Using four polymorphic allozymes, we compared the outcrossing-rate estimates of two groups of individuals under natural conditions: one group with low herkogamy and another with high herkogamy. The high herkogamy group had a higher outcrossing rate (0.572) than the low herkogamy group (0.335). This suggests that the within-population variation in outcrossing rate could potentially cause the previously observed association between herkogamy and inbreeding depression (Takebayashi and Delph, 2000). A previous study of floral traits among G. achilleifolia populations failed to detect a relationship between herkogamy and outcrossing rate, demonstrating that the functionality of traits may be obscured in among-population studies as a consequence of uncontrolled environmental variation. Additionally, the effect of herkogamy on outcrossing rate in delayed selfers such as G. achilleifolia may be particularly prominent when pollinator availability is low. Our population-level estimate of outcrossing rate (0.444) was somewhat lower than an estimate from the same population, 15 years prior to our study (0.75), suggesting that pollinator availability may fluctuate among years. Both within-year and among-year variation in the outcrossing rate may have a strong influence on mating-system evolution.

Analysis of Variance↗

Host and parasite population structure in a natural plant-pathogen system

We investigated the genetic population structure in a metapopulation of the plant Silene latifolia (Caryophyllaceae) and its fungal pathogen Microbotryum violaceum (Ustilaginales), a pollinator-borne disease. Population structure of the host plant was estimated using allozyme markers and that of the fungus by microsatellites. Both host and parasite showed significant differentiation, but parasite populations were 12 times more strongly differentiated than those of the hosts. We found significant isolation by distance for host populations but not for parasite populations. Higher population differentiation for the parasite may result from small effective population size, high selfing rates, or low migration rate. In this system, hosts are obligate outcrossers and they migrate by seeds and pollen, whereas parasites can self-fertilize and migrate only on pollinating insects. We discuss the effect of limited gene flow in this parasite on its coevolutionary interaction with its host, and its potential for local adaptation on sympatric host populations.

Journal Article↗

Regulation of triacylglycerol biosynthesis in embryos and microsomal preparations from the developing seeds of Cuphea lanceolata.

Embryos of Cuphea lanceolata have more than 80 mol% of decanoic acid ('capric acid') in their triacylglycerols, while this fatty acid is virtually absent in phosphatidylcholine (PtdCho). Seed development was complete 25-27 days after pollination, with rapid triacylglycerol deposition occurring between 9 and 24 days. PtdCho amounts increased until day 15 after pollination. Analysis of embryo lipids showed that the diacylglycerol (DAG) pool consisted of mainly long-chain molecular species, with a very small amount of mixed medium-chain/long-chain glycerols. Almost 100% of the fatty acid at position sn-2 in triacylglycerols (TAG) was decanoic acid. When equimolar mixtures of [14C]decanoic and [14C]oleic acid were fed to whole detached embryos, over half of the radioactivity in the DAG resided in [14C]oleate, whereas [14C]decanoic acid accounted for 93% of the label in the TAG. Microsomal preparations from developing embryos at the mid-stage of TAG accumulation catalysed the acylation of [14C]glycerol 3-phosphate with either decanoyl-CoA or oleoyl-CoA, resulting in the formation of phosphatidic acid (PtdOH), DAG and TAG. Very little [14C]glycerol entered PtdCho. In combined incubations, with an equimolar supply of [14C]oleoyl-CoA and [14C]decanoyl-CoA in the presence of glycerol 3-phosphate, the synthesized PtdCho species consisted to 95% of didecanoic and dioleic species. The didecanoyl-glycerols were very selectively utilized over the dioleoylglycerols in the production of TAG. Substantial amounts of [14C]oleate, but not [14C]decanoate, entered PtdCho. The microsomal preparations of developing embryos were used to assess the acyl specificities of the acyl-CoA:sn-glycerol-3-phosphate acyltransferase (GPAT, EC 2.3.1.15) and the acyl-CoA:sn-1-acyl-glycerol-3-phosphate acyltransferase (LPAAT, EC 2.3.1.51) in Cuphea lanceolata embryos. The efficiency of acyl-CoA utilization by the GPAT was in the order decanoyl = dodecanoyl greater than linoleoyl greater than myristoyl = oleoyl greater than palmitoyl. Decanoyl-CoA was the only acyl donor to be utilized to any extent by the LPAAT when sn-decanoylglycerol 3-phosphate was the acyl acceptor. sn-1-Acylglycerol 3-phosphates with acyl groups shorter than 16 carbon atoms did not serve as acyl acceptors for long-chain (greater than or equal to 16 carbon atoms) acyl-CoA species. On the basis of the results obtained, we propose a schematic model for triacylglycerol assembly and PtdCho synthesis in a tissue specialized in the synthesis of high amounts of medium-chain fatty acids.

Acyl Coenzyme A↗

Decrease of allergen-specific T-cell response induced by local nasal immunotherapy.

BACKGROUND: The clinical efficacy and safety of local nasal immunotherapy (LNIT) with lyophilized 'macronized' powder has been demonstrated. However, the immunological changes possibly induced by LNIT which may account for the clinical improvement are still unclear. OBJECTIVE: To investigate the effects of a successful LNIT-treatment on the allergen-driven T cell response, cytokine secretion and IgE and IgG antibody production. METHODS: Three groups (untreated, subcutaneous immunotherapy- SIT- and LNIT-treated) of grass-sensitive patients suffering from seasonal rhinitic symptoms were ramdomized for the 2-year study. The proliferative response of PBMC to purified Rye-1 allergen and serum levels of grass-specific IgE and IgG were evaluated before treatment and during the 2-year subsequent pollination periods. The proliferative response of allergen-specific short-term T-cell lines, as well as production of allergen-driven cytokine by PBMC, were also assessed. RESULTS: Both SIT and LNIT induced a significant reduction of symptom scores during the pollination season. SIT, but not LNIT, induced a significant change in serum levels of allergen-specific IgE and IgG antibody. By contrast, both SIT and LNIT reduced the increase of the proliferative response of allergen-specific T cells driven by natural allergen exposure and significantly decreased T cell proliferation to low doses of allergen, as shown also by the mitogenic index of allergen-specific T-cell lines. A reduced IL-4 and IFNgamma production by PBMC of LNIT- and SIT-treated patients was also observed in the absence of a clearcut TH2-TH1 switch. CONCLUSIONS: These data suggest that a common mechanism of both LNIT and SIT is the induction of T-cell tolerance, thus providing a rational basis to explain why LNIT may be clinically successful in allergic patients with rhinits.

Administration, Intranasal↗

Evolution and maintenance of stigma-height dimorphism in Narcissus. II. Fitness comparisons between style morphs.

Populations of the insect-pollinated geophytes Narcissus assoanus and N. dubius (Amaryllidaceae) are commonly dimorphic for stigma height. An extensive survey of populations of the two species in SW France revealed a wide range of style-morph frequencies, particularly populations with significantly more long-styled than short-styled plants. Here we employ experimental and theoretical approaches to investigate potential selective mechanisms governing the variation in style-morph frequencies. Controlled pollination of both species demonstrated that N. assoanus is moderately self-sterile whereas N. dubius is highly self-compatible. Both intra- and intermorph crosses of N. assoanus were equally fertile, indicating that the species does not exhibit heteromorphic incompatibility. Estimates of female fertility (fruit- and seed-set) and multilocus estimates of outcrossing using allozyme markers provided no evidence of morph-specific differences in maternal components of reproductive success in natural populations of the two species. This result suggested that differences between the morphs in male fertility may be largely responsible for the observed morph-ratio variation. To investigate this hypothesis we developed a mating model that incorporates the genetics of stigma-height dimorphism and contrasting rates of assortative and disassortative mating in the style morphs. Simulation results demonstrated that stigma-height dimorphism will always be maintained when levels of disassortative mating are greater than assortative mating, and that the observed L-biased populations in Narcissus spp. probably result from greater levels of assortative mating in this morph in comparison with the S-morph.

Biological Evolution↗

F1 hybrid inviability in eucalyptus: the case of E. ovata x E. globulus.

The impact of inbreeding and hybridization on fitness was compared in the two co-occurring forest tree species, Eucalyptus ovata and E. globulus, aimed at explaining the rarity of their hybrids in nature. The success of selfing, open-pollination and outcrossing of both species and interspecific hybridization was monitored from seed-set to 10-year's growth in a field trial. There was a unilateral barrier to hybridization with seed-set obtained only with E. ovata females. The F1 hybrids exhibited reduced viability compared to intraspecific cross-types at virtually all stages of the life cycle and are clearly at a selective disadvantage compared with their open-pollinated E. ovata half-sibs with which they would directly compete in nature. Eucalyptus ovata and E. globulus overlap in their flowering time but the F1 hybrids flowered later with virtually no overlap with either species. The asynchronous flowering and reduced reproductive fitness of F1 hybrids would markedly limit the opportunity for advanced generation hybridization. Inbreeding similarly had a deleterious effect on the fitness of both species, and the F1 hybrids were most competitive with the E. ovata selfs. It is argued that changes in inbreeding levels of parental populations may be a key factor affecting the relative fitness of hybrids and their potential to impact on the pure species gene pool. Reduced fitness of the pure species through inbreeding may result in hybridization having its greatest evolutionary impact in small founder or relict populations.

Biological Evolution↗

Mating system parameters of Dryobalanops aromatica Gaertn. f. (Dipterocarpaceae) in three different forest types and a seed orchard.

The mating system of Dryobalanops aromatica in three different forest types and a seed orchard was quantified by allozyme analysis of progeny arrays using a mixed-mating model. The primary forest (Bukit Sai) had the highest multilocus outcrossing rate (tm=0.923 +/- 0.035), followed by logged forest (Lesong; tm=0.766 +/- 0.056) and artificial forest (FRIM; tm=0.661 +/- 0.066) with seed orchard showing the lowest (Tampin; tm=0.551 +/- 0.095). Deviations from the mixed mating model were evident from differences in pollen and ovule allele frequencies, and heterogeneity of pollen pools in all three different forest types and the seed orchard. A high rate of outcrossing in primary forest (tm=0.92) may indicate that the species is self-incompatible, but a lower value in the seed orchard (tm=0.55) might suggest further that the self-incompatibility system is weak. The outcrossing rate was greater in the primary forest (tm=0.92) than in logged forest (tm=0.77). It is argued that this might be a consequence of the lower density of flowering trees and alteration of pollinator foraging behaviour in logged forest. Higher values of correlated mating (rp) and biparental mating (tm - ts) in primary forest (0.08 and 0.39, respectively) in comparison with logged forest (0.03 and 0.11, respectively) demonstrate that logging activities might reduce the seeds produced through consanguineous mating. Compared with primary forest, it is argued that lower rates of outcrossing in artificial forest (tm=0.67) and seed orchard (tm=0. 55) might be attributed to lack of flowering synchrony and insufficient number of pollinators. The high level of correlated mating (rp=0.43) and biparental mating (tm - ts=0.12) in the seed orchard may further suggest that the seed orchard was established using related seed sources.

Crosses, Genetic↗

Genetic variation within and between populations of Pinus sylvestris L. (Scots pine) for susceptibility to Melampsora pinitorqua Rostr. (pine twist rust).

The genetic variability of Scots pine (Pinus sylvestris L.) for twist rust susceptibility and the consequences for genetic improvement of the species were evaluated after artificial inoculation of 1-year-old seedlings in greenhouse tests. Wind-pollinated progenies, factorial and incomplete diallel mating designs were used to compare two natural populations of Scots pine (Haguenau, France, and Taborz, Poland) and their hybrids (Haguenau x Taborz). Families from Taborz were significantly (P < 0.001) less susceptible to twist rust than those from Haguenau. Inter-population hybrids were as susceptible as the Haguenau population. No clear relationship between severity of infection, shoot length and phenological stage at time of inoculation was observed in intra- and interpopulation crosses. Differences observed between the two natural populations for their response to twist rust are suggested to be the consequence of local adaptation simultaneously for climatic conditions and pathogen pressure. Within each of the two populations, the effects of general combining abilities (GCA) for rust susceptibility were predominant, although estimation of individual heritabilities remained at a moderate level (0.30-0.54). In interpopulation crosses, GCA effects of Haguenau and Taborz populations were still predominant. Moreover, hybrid performance could be predicted with confidence based on the intrapopulation GCA values of parents. Genetic gain on twist rust resistance could be achieved quickly through intrapopulation mass or backward selection based on wind-pollinated progeny tests.

Basidiomycota↗

Inbreeding depression in two populations of Arenaria uniflora (Caryophyllaceae) with contrasting mating systems.

I used parallel family-structured crossing designs to investigate the relative performance of self and outcross progeny in selfing and predominantly outcrossing populations of the annual plant Arenaria uniflora. The selfer population experienced much lower inbreeding depression (delta = 0.05 +/- 0.02 SE) than the outcrossers (delta = 0.19 +/- 0.02 SE). The negative association between genetic load and selfing rate suggests that purgable partially recessive alleles are the primary source of inbreeding depression, as does its late expression in both populations. Inbreeding depression in the selfer population, which naturally consists of highly inbred lines, was used to calculate the mean dominance (h = 0.33) and incidence rate (U = 0.30) of deleterious mutations. In the outcrosser population, significant variation among individuals in the expression of inbreeding depression may reflect lineage-specific differences in inbreeding history or, more probably, random variation in mutational load. The low (<< 0.5) inbreeding depression of outcrossers suggests that the maintenance of a mixed mating system in some A. uniflora populations and the evolution of nearly cleistogamous self-pollination in others may reflect local pollinator-mediated selection for selfing rather than the constant 3:2 genetic advantage invoked by many models.

Analysis of Variance↗

Low neighbourhood size and high interpopulation differentiation in the endangered shrub Grevillea iaspicula McGill (Proteaceae).

Mating system parameters and genetic diversity were examined for five populations of the endangered shrub Grevillea iaspicula (Proteaceae). Controlled pollinations show that G. iaspicula has an effective self-incompatibility system and little potential for agamospermy. This is reflected in uniformly high multilocus outcrossing rates (tm=0.96-1.00). However, average paternal diversity within open-pollinated sibships is low (rp=0.31-0.54), suggesting that mating within populations is quite restricted. Despite the small size of most populations (four of the five populations studied have fewer than 20 reproductive individuals) the species still possesses moderate to high allelic richness (A=1.6-2.5). Interpopulation genetic differentiation is high (D=0.04-0.32), suggesting that gene flow is limited, even among populations separated by only a few kilometres.

Conservation of Natural Resources↗