The effects of plant density upon pollination success, reproductive effort and fruit parasitism in Cistus ladanifer L. (Cistaceae)

Human activity often causes a decline in the local density of plant populations. Below some critical lower density, populations may suffer a progressive decline in reproductive success because of the difficulties associated with finding suitable mates. Therefore, to conserve endangered plant species it is necessary to understand in greater detail how changes in population density affect different determinants of plant reproductive success. We simultaneously recorded individual plant pollination success, reproductive effort and fruit parasitism in three populations of Cistus ladanifer L. in eastern Portugal. Pollination success declined significantly as distance to the nearest conspecific increased (p<0.001). However, reproductive effort and fruit parasitism showed the opposite pattern (both p<0.001). On average, plants farther than 1 m from their nearest conspecific suffered from three times more fruit parasitism compared to plants closer to a conspecific. Overall, net female reproductive output decreased as nearest neighbor distance rose (p<0.001). Thus, isolated plants were able to compensate only partially for reduced pollination success through increased reproductive effort. We conclude that management plans for plant populations should recognize that reproductive success is the accumulated result of several different processes, which may each respond to plant density in different ways.     

The role of individual variation in flowering and pollination in the reproductive success of a crepuscular buzz-pollinated plant

Background and AimsPlant individuals within a population differ in their phenology and interactions with pollinators. However, it is still unknown how individual differences affect the reproductive success of plants that have functionally specialized pollination systems. Here, we evaluated whether plant individual specialization in phenology (temporal specialization) and in pollination (pollinator specialization) affect the reproductive success of the crepuscular-bee-pollinated plant Trembleya laniflora (Melastomataceae).MethodsWe quantified flowering activity (amplitude, duration and overlap), plant–pollinator interactions (number of flowers visited by pollinators) and reproductive success (fruit set) of T. laniflora individuals from three distinct locations in rupestrian grasslands of southeastern Brazil. We estimated the degree of individual temporal specialization in flowering phenology and of individual specialization in plant–pollinator interactions, and tested their relationship with plant reproductive success.Key Results Trembleya laniflora presented overlapping flowering, a temporal generalization and specialized pollinator interactions. Flowering overlap among individuals and populations was higher than expected by chance but did not affect the individual interactions with pollinators and nor their reproductive success. In contrast, higher individual generalization in the interactions with pollinators was related to higher individual reproductive success.ConclusionsOur findings suggest that individual generalization in plant–pollinator interaction reduces the potential costs of specialization at the species level, ensuring reproductive success. Altogether, our results highlight the complexity of specialization/generalization of plant–pollinator interactions at distinct levels of organization, from individuals to populations, to species.     

The effect of isozyme selection on metric characters in an interspecific backcross of tomato — basis of an early screening procedure

Summary The extent of correlation was estimated between isozyme genotypes and the four widely segregating characters — leaf segment W/L ratio, stigma exsertion, fruit weight, and seed weight — in the first backcross of F₁ Lycopersicon esculentum x Solanum pennellii to the former parent. The inbred parents differ in their alleles at the 12 tested isozymic loci, which are known to mark a minimum of eight of the twelve tomato chromosomes. Based on the isozyme data, a mean heterozygosity value, ¯H, was calculated which estimates the proportion of pennillii alleles in each individual. Correlations between mean heterozygosity and observed levels of each quantitative trait were highly significant and positive or negative as expected from the relative parental values. Plants with the lowest mean heterozygosity — i.e., closest to the esculentum zymotype also had mean values closest to those of this parent amongst the whole backcross population for each of the quantitative traits.Bivariate and multiple regression analysis was used to evaluate the ability of isozymes vs diagnostic morphological characters to estimate the portion of recurrent parent genes carried in each backcross individual. The results suggest that isozyme data gives better estimates than single diagnostic morphological characters and approach the level obtained by combinations of three morphological traits. Since electrophoretic determinations are made on small seedlings, selection at that stage can effect great savings of space and effort by greatly deminishing the size of the population needed at maturity. As such, isozyme selection would precede morphological selection but not replace it, thus the predictive value of these biochemical markers as well as diagnostic morphological characters could be obtained.     

Effects of plant size and weather on the flowering phenology of the organ pipe cactus (Stenocereus thurberi)

Background and Aims

Flowering phenology is a critical life-history trait that influences reproductive success. It has been shown that genetic, climatic and other factors such as plant size affect the timing of flowering and its duration. The spatial and temporal variation in the reproductive phenology of the columnar cactus Stenocereus thurberi and its association with plant size and environmental cues was studied.

Methods

Flowering was monitored during 3 years in three populations of S. thurberi along a latitudinal gradient. Plant size was related to phenological parameters. The actual and past weather were used for each site and year to investigate the environmental correlates of flowering.

Key Results

There was significant variation in the timing of flowering within and among populations. Flowering lasted 4 months in the southern population and only 2 months in the northern population. A single flowering peak was evident in each population, but ocurred at different times. Large plants produced more flowers, and bloomed earlier and for a longer period than small plants. Population synchrony increased as the mean duration of flowering per individual decreased. The onset of flowering is primarily related to the variance in winter minimum temperatures and the duration to the autumn–winter mean maximum temperature, whereas spring mean maximum temperature is best correlated with synchrony.

Conclusions

Plant size affects individual plant fecundity as well as flowering time. Thus the population structure strongly affects flowering phenology. Indications of clinal variation in the timing of flowering and reproductive effort suggest selection pressures related to the arrival of migrating pollinators, climate and resource economy in a desert environment. These pressures are likely to be relaxed in populations where individual plants can attain large sizes.Key words: Flowering phenology, optimal timing, plant size, Sonoran Desert, Stenocereus thurberi, temperature     

Phenotypic Plasticity and Population Differentiation in an Ongoing Species Invasion

The ability to succeed in diverse conditions is a key factor allowing introduced species to successfully invade and spread across new areas. Two non-exclusive factors have been suggested to promote this ability: adaptive phenotypic plasticity of individuals, and the evolution of locally adapted populations in the new range. We investigated these individual and population-level factors in Polygonum cespitosum, an Asian annual that has recently become invasive in northeastern North America. We characterized individual fitness, life-history, and functional plasticity in response to two contrasting glasshouse habitat treatments (full sun/dry soil and understory shade/moist soil) in 165 genotypes sampled from nine geographically separate populations representing the range of light and soil moisture conditions the species inhabits in this region. Polygonum cespitosum genotypes from these introduced-range populations expressed broadly similar plasticity patterns. In response to full sun, dry conditions, genotypes from all populations increased photosynthetic rate, water use efficiency, and allocation to root tissues, dramatically increasing reproductive fitness compared to phenotypes expressed in simulated understory shade. Although there were subtle among-population differences in mean trait values as well as in the slope of plastic responses, these population differences did not reflect local adaptation to environmental conditions measured at the population sites of origin. Instead, certain populations expressed higher fitness in both glasshouse habitat treatments. We also compared the introduced-range populations to a single population from the native Asian range, and found that the native population had delayed phenology, limited functional plasticity, and lower fitness in both experimental environments compared with the introduced-range populations. Our results indicate that the future spread of P. cespitosum in its introduced range will likely be fueled by populations consisting of individuals able to express high fitness across diverse light and moisture conditions, rather than by the evolution of locally specialized populations.     

Assessment of allozyme variation among New Zealand populations of Gracilaria chilensis (Gracialiares,Rhodophyta) using starch-gel electrophoresis

New Zealand populations of Gracilaria chilensis are uniform in anatomical reproductive characteristics but vary morphologically and have been found to separate into two distinct groups with respect to agar methylation level, namely low (24–30%) and high (43–47%). To investigate the genetic variation within New Zealand populations of this species, 14 isozyme loci detected by starch-gel electrophoresis were examined in 17 wild populations from a wide range of localities, and in cultures derived from these populations. Five of these loci were polymorphic, but the genetic variation within populations was low: of the 17 populations examined, 15 were fixed at all loci (heterozygosity 0.000) and in the remaining two populations the observed heterozygosity was still low (0.004 and 0.011). The genetic distances between the populations ranged from 0.00 to 0.43. UPGMA cluster analysis separated the populations into two groups, a northern group and a group found throughout the country. Although these two groups do not correlate with the two groups based on agar methylation level at every locality, the correlation is sufficiently striking to merit further investigation.     

Heterosis at Allozyme Loci under Inbreeding and Crossbreeding in PINUS ATTENUATA

The dependence of heterosis at isozyme loci on inbreeding and crossbreeding was studied in 10-yr-old trees of knobcone pine (Pinus attenuata Lemm.). Heterozygosity was determined at 24 polymorphic isozyme loci and related to the rate of vegetative growth and cone production. The inbreds, created by selfpollination, had 46% of the heterozygosity of their mothers; the crossbreds, created by interpopulation crossing, had 155% of the heterozygosity of their mothers. Within the crossbreds, heterozygosity was positively correlated with trunk growth, but negatively correlated with cone production. Results in the crossbreds, however, were strongly influenced by a few individuals that showed unusually slow growth, high reproduction and low heterozygosity. Without those individuals, there was no relationship of heterozygosity to either growth or reproduction.—Within the inbreds, heterozygosity was positively correlated with both trunk growth and cone production. Each locus that was heterozygous in the mothers was calculated to mark about 3% of the genome for identity by descent in the inbred progeny; the total proportion of the genome marked was between 10 and 11%. Using these estimates to relate heterozygosity to the inbreeding coefficient (F) gave estimates of inbreeding depression per unit of F that fell within the range of published values for conifers. The strength of heterosis found among the inbreds suggests that single-locus or multilocus overdominance should be exceedingly difficult to detect in natural populations of predominantly outcrossing species.     

Isozyme characterization of genetic diversity in Minnesota populations of purple loosestrife,Lythrum salicaria (lythraceae)

Starch gel electrophoresis of plant proteins was used to identify purple loosestrife (Lythrum spp.) cultivars and weedy populations. Preliminary determinations were made as to what degree weedy loosestrife populations were related (or genetically similar) to populations of L. alatum, L. virgatum, and horticultural cultivars. Cluster analysis of the data indicated that native L. alatum was genetically different from all populations of purple loosestrife and cultivars examined. The L. salicaria and L. virgatum cultivars, as groups, were not genetically distinguishable from the weedy populations analyzed. Seven cultivars of L. salicaria origin analyzed as a group were not distinguishable from the eight cultivars of L. virgatum origin, indicating that separation by cultivar origin may not be feasible. While the two “groups” were not distinguishable, most individual cultivars could be distinguished from one another by isozyme phenotype. Genetic variation was high within populations of weedy purple loosestrife but low among populations, which is characteristic of polyploid, perennial plant species that are widely distributed. Geographic location did not consistently correlate with genetic similarity.     

Genetic Diversity and Population Structure in Polygonum cespitosum: Insights to an Ongoing Plant Invasion

Molecular markers can help elucidate how neutral evolutionary forces and introduction history contribute to genetic variation in invaders. We examined genetic diversity, population structure and colonization patterns in the invasive Polygonum cespitosum, a highly selfing, tetraploid Asian annual introduced to North America. We used nine diploidized polymorphic microsatellite markers to study 16 populations in the introduced range (northeastern North America), via the analyses of 516 individuals, and asked the following questions: 1) Do populations have differing levels of within-population genetic diversity? 2) Do populations form distinct genetic clusters? 3) Does population structure reflect either geographic distances or habitat similarities? We found low heterozygosity in all populations, consistent with the selfing mating system of P. cespitosum. Despite the high selfing levels, we found substantial genetic variation within and among P. cespitosum populations, based on the percentage of polymorphic loci, allelic richness, and expected heterozygosity. Inferences from individual assignment tests (Bayesian clustering) and pairwise F _ST values indicated high among-population differentiation, which indicates that the effects of gene flow are limited relative to those of genetic drift, probably due to the high selfing rates and the limited seed dispersal ability of P. cespitosum. Population structure did not reflect a pattern of isolation by distance nor was it related to habitat similarities. Rather, population structure appears to be the result of the random movement of propagules across the introduced range, possibly associated with human dispersal. Furthermore, the high population differentiation, genetic diversity, and fine-scale genetic structure (populations founded by individuals from different genetic sources) in the introduced range suggest that multiple introductions to this region may have occurred. High genetic diversity may further contribute to the invasive success of P. cespitosum in its introduced range.     

Genetic Variation and Differentiation of Peat-Bog and Dry-Meadow Populations of the Dwarf Mountain Pine Pinus mugo Turra in the Highlands of the Ukrainian Carpathians

Based on electrophoretic analysis of 21 isozyme loci controlling 10 enzyme systems, the intra- and interpopulation variation was studied in two peat-bog and three dry-meadow populations of the dwarf mountain pine Pinus mugoTurra from the highlands of the Ukrainian Carpathians. In the studied samples (a total of 164 trees), on average 62% of the studied genes were polymorphic; the mean heterozygosity was 21.3%. The dry-meadow populations differed from the peat-bog populations by allele and genotype diversity and by heterozygosity although the indices characterizing population heterogeneity (F _st and G _st) were small (0.027 and 0.032, respectively). Nei's genetic distances between the populations ranged of 0.011 to 0.032 with the mean of 0.018.     

Inbreeding depression influences genet size distribution in a marine angiosperm

Although inbreeding depression is a major genetic phenomena influencing individual fitness, it is difficult to measure in wild populations. An alternative approach is to correlate heterozygosity, measured using highly polymorphic markers, with a fitness-correlated trait. In clonal plants, genet size is predicted to be fitness correlated. Here we test the prediction that the genet size distribution of the marine clonal plant Zostera marina (eelgrass) is influenced by inbreeding depression. We used nine polymorphic microsatellite markers to access the fine scale clonal structure and to measure individual heterozygosity within 4 plots (each corresponds to 256 m2, sampled at 1-m intervals) in two populations along the German Baltic Coast. The same plots were also sampled for flowering and vegetative shoots to obtain estimates for sexual reproductive output at the level of the genetic individual. We found substantial differences in the genet size distribution between the two populations that may be explained by different disturbance frequency. In both populations, clone size was significantly positively correlated with the total number of flowering shoots, indicating that larger clones have a higher reproductive output. Individual heterozygosity was significantly positively associated with clone size. The effect was much stronger in Falkenstein (low disturbance) than in Maasholm (high disturbance). The results indicate that in a low disturbance population the relatively outbred clones occupy a higher proportion of the available space, possibly because they outcompete relatively inbred neighbours.     

Reproductive allocation strategies in desert and Mediterranean populations of annual plants grown with and without water stress

Reproductive effort (relative allocation of biomass to diaspore production) was compared in matched pairs of Mediterranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Bromus fasciculatus C. Presl. and Brachypodium distachyon (L.) Beauv., grown under high and low levels of water availability in a common-environment experiment. Desert populations in all three species showed higher reproductive effort than corresponding Mediterranean populations, as expressed by both a reproductive index (RI= reproductive biomass/vegetative biomass), and a reproductive efficiency index (REI=number of diaspores/total plant biomass). Moreover, in E. hispanica and Brachypodium distachyon, inter-populational differences in reproductive effort were greater under water stress, the main limiting factor for plant growth in the desert. These results indicate that variability in reproductive effort in response to drought is a critical and dynamic component of life history strategies in annual species in heterogeneous, unpredictable xeric environments. When subjected to water stress the Mediterranean populations of E. hispanica and B. distachyon showed greater plasticity (e.g. had a greater reduction) in reproductive effort than the desert populations, while in Bromus fasciculatus both populations showed similar amounts of plasticity.     

Genetic diversity of nine faba bean (Vicia faba L.) populations revealed by isozyme markers

Seven isozyme systems (Sod, 6-Pgd, Me, Est, Skdh, Fdh and Gdh) representing nine loci were used to study the genetic diversity of nine faba bean populations. Seven loci revealed polymorphic bands and showed the same quaternary structure as that found in several species. They revealed a high number of phenotypes. Indeed, from 3 to 9 phenotypes per locus were investigated in this study. The percentage of polymorphic loci (P = 59.3 %) was higher than that mentioned in the autogamous species (P = 20.3 %) and less than the optimum (P=96 %) indicated for allogamous plants. Total genetic diversity (H _T) and within population genetic diversity (H _S) were estimated with the isozyme markers. The contribution of among population genetic diversity (D _ST) to total genetic diversity was 22%. Enzyme markers pointed out an average inbreeding level for whole population (F _IT) and within population (F _IS). Within population genetic diversity represents 78% of total diversity. Intra-population genetic diversity (H _S = 0.206) was ranged with the respect of allogamous species and was clearly higher than that of among population genetic diversity (D _ST = 0.057) indicating an out-crossing predominance in the studied populations. The expected heterozygosity was higher than that observed heterozygosity at the allogamous species was confirmed in this study. Although, the mean estimated gene flow was less than 1(Nm=0.814), the dendrogram based on Nei’s genetic distance of the 9 populations using UPGMA method showed some genetic drift between populations.     

COMPONENTS OF REPRODUCTIVE EFFORT AND YIELD IN GOLDENRODS

Four components of reproductive yield (the weight of reproductive tissue) were examined in relation to their effect on reproductive effort and their relative contributions to reproductive yield in five species of goldenrods (Solidago, Compositae). The yield components were number of flowing stems per plant, number of flowering branches per stem, number of flowering heads per branch, and number of seeds per seed head. Individuals within populations increase their reproductive effort by increasing their reproductive weight, not by decreasing their vegetative weight. Each species shows a different pattern of positive correlations of yield components with reproductive yield and reproductive effort, indicating that each species has its own mechanisms for regulating reproduction. The yield components were not significantly intercorrelated.     

Genetic variation relevant for the conservation of the narrow endemic Senecio carbonensis (Asteraceae) from the southern Andes

We studied morphological variation, reproductive biology and genetic structure of Senecio carbonensis, a narrow endemic from high elevations of the southern Andes, and compared it to Senecio peteroanus, a closely related widespread congeneric species. Using ANOVAs and discriminant analysis we showed that populations of S. carbonensis were more similar to each other in their morphology, had comparatively limited reproductive capacity and presented lower plant density and more reduced plant cover than populations of S. peteroanus. Similar high genetic variation was found at the population and species level in both species, based on isozyme variation at 14 and 11 putative gene loci that were resolved in S. carbonensis and S. peteroanus, respectively. The two species were genetically distinct. However, the small genetic distance between populations of each species suggests that either they were recently founded and that genetic drift did not have time to promote divergence, or that its effects are masked by significant current gene flow. These results show that ecological and reproductive characteristics of S. carbonensis may limit its extension and abundance. In contrast, high genetic variation at the population level is probably favored by self‐incompatibility. Despite the restricted geographical distribution and low plant density of S. carbonensis populations, genetic characteristics do not seem to be limiting their long‐term persistence. This information suggests that populations of S. carbonensis are not at risk of extinction as a result of genetic factors, although it is important to study and monitor population dynamics to further assess the degree of recruitment through time.     

Isozyme diversity and genetic structure of buckwheat populations

Genetic diversity of 35 Fagopyrum populations belonging to thirteen taxa was studied using isozyme variation at 16 loci for nine enzyme systems (ADH, α-AMY, EST, GDH, GOT, IDH, MDH, PGM, and POD). The mean number of alleles per locus (A_p) showed little variation, ranging from 1.38 to 1.71. The mean percentage of polymorphic (P) of the thirteen taxa was high, ranging from 68.75% to 93.75. The mean expected heterozygosity (H_e) ranged from 0.21 to 0.37. Obvious grouping of populations according to taxonomic species was found in the cluster analysis, confirmed that the thirteen taxa studied were independent, with a clear boundary. Based on the dendrogram, the phylogenic relationship of Fagopyrum species are in agreement with the achene size were found, and it was inferred that southwestern China should be the original center of cultivated buckwheat.     

Genetic relationships among twelve Chinese indigenous goat populations based on microsatellite analysis

Twelve Chinese indigenous goat populations were genotyped for twenty-six microsatellite markers recommended by the EU Sheep and Goat Biodiversity Project. A total of 452 goats were tested. Seventeen of the 26 microsatellite markers used in this analysis had four or more alleles. The mean expected heterozygosity and the mean observed heterozygosity for the population varied from 0.611 to 0.784 and 0.602 to 0.783 respectively. The mean F_ST (0.105) demonstrated that about 89.5% of the total genetic variation was due to the genetic differentiation within each population. A phylogenetic tree based on the Nei (1978) standard genetic distance displayed a remarkable degree of consistency with their different geographical origins and their presumed migration throughout China. The correspondence analysis did not only distinguish population groups, but also confirmed the above results, classifying the important populations contributing to diversity. Additionally, some specific alleles were shown to be important in the construction of the population structure. The study analyzed the recent origins of these populations and contributed to the knowledge and genetic characterization of Chinese indigenous goat populations. In addition, the seventeen microsatellites recommended by the EU Sheep and Goat Biodiversity Project proved to be useful for the biodiversity studies in goat breeds.     

Relationships between heterozygosity and fitness in the Iberian threatened larkspur Delphinium bolosii (Ranunculaceae)

The relationship between individual heretozygosity and fitness was explored in the perennial larkspur Delphinium bolosii (Ranunculaceae), an endangered species endemic to Catalonia (North-Eastern Spain), as a part of several studies prior to designing a programme for the reintroduction of this species in a locality where it has been extinct for approximately one century. Allozyme electrophoresis was used to quantify the levels of heterozygosity at nearly to one hundred surveyed individuals in two extant populations. At the same time, eight parameters of vegetative and reproductive fitness were measured. A principal component analysis reduced the original fitness variables to three uncorrelated principal components, two associated with the maternal plant and the other one with the offspring. However, none of the components were significantly correlated with heterozygosity. The low number of variable allozyme markers and the likely influence of ecological factors could be responsible for the lack of correlation between individual heterozygosity and fitness.     

Herbivore host-associated genetic differentiation depends on the scale of plant genetic variation examined

Herbivore adaptation to plant genetic variation can lead to reproductive isolation and the formation of host-associated lineages (host-associated differentiation, or HAD). Plant genetic variation exists along a scale, ranging from variation among individual plant genotypes to variation among plant species. Along this scale, herbivores may adapt and diverge at any level, yet few studies have examined whether herbivore differentiation exhibits scaling with respect to host variation (e.g., from genotypes to species). Determining at which level(s) herbivore differentiation occurs can provide insight into the importance of plant genetic variation on herbivore evolution. Previous studies have found strong genetic differentiation in the eriophyid mite, Aceria parapopuli, between hybrid Populus hosts and parental Populus species, but minimal neutral-locus differentiation among individual trees of the same species. We tested whether genetic differentiation in A. parapopuli scales with genetic variation in its Populus hosts. Using mite ITS1 sequence data collected among host species and among host populations, two key patterns emerged. (1) We found strong differentiation of A. parapopuli among Populus species, supporting the hypothesis that plant species differences drive reproductive isolation and HAD. (2) We did not find evidence of host-driven genetic differentiation in mites at the level of plant populations, suggesting that this level of plant variation is insufficiently strong to drive differentiation at a neutral locus. In combination with previous studies, we found that HAD occurs at the higher levels of plant genetic variation, but not at lower levels, and conclude that HAD depends on the scale of plant genetic variation examined.     

Within and between species variation in response to environmental gradients in Polygonum pensylvanicum and Polygonum virginianum

Summary The responses of Polygonum pensylvanicum L., an early successional annual, and of Polygonum virginianum L., a late successional perennial, were examined along discontinuous gradients of soil moisture, light and nutrient availability. In the case of P. virginianum both individuals grown from seed and individuals grown from rhizomes were examined. The results show that variation in the response of individuals of a species of different age to environmental variation is as great as that found between the two congeneric species of different successional habitats. In general, individuals of the two species were more similar to one another in response to the resource gradients when both were started from seed, than were individuals of P. virginianum grown from seed and from rhizomes. Potential differences in stored reserves (starting capital) between rhizomes and seeds appeared to have little effect upon responses to resource availability. A number of plant characters were found to vary along the gradients in ways that were unique to the character, the gradient, and the age of the individual. These characters included aspects of leaf size, shape, and orientation, as well as whole plant architecture. Many if not all of these characters are likely to have significant effects upon the functioning of plants. The origin of this difference in response to the gradients of individuals of P. virginianum of differing age may be ontogenetic or may reflect differences in genetic composition created by recombination.