Late-flowering Populations Research Articles

Identification of Differentially Expressed Genes Related to Floral Bud Differentiation and Flowering Time in Three Populations of Lycoris radiata.

The transition from vegetative to reproductive growth is important for controlling the flowering of Lycoris radiata. However, the genetic control of this complex developmental process remains unclear. In this study, 18 shoot apical meristem (SAM) samples were collected from early-, mid- and late-flowering populations during floral bud differentiation. The histological analysis of paraffin sections showed that the floral bud differentiation could be divided into six stages; the differentiation time of the early group was earlier than that of the middle and late groups, and the late group was the latest. In different populations, some important differential genes affecting the flowering time were identified by transcriptome profiles of floral bud differentiation samples. Weighted gene co-expression network analysis (WGCNA) was performed to enrich the gene co-expression modules of diverse flowering time populations (FT) and floral bud differentiation stages (ST). In the MEyellow module, five core hub genes were identified, including CO14, GI, SPL8, SPL9, and SPL15. The correlation network of hub genes showed that they interact with SPLs, AP2, hormone response factors (auxin, gibberellin, ethylene, and abscisic acid), and several transcription factors (MADS-box transcription factor, bHLH, MYB, and NAC3). It suggests the important role of these genes and the complex molecular mechanism of floral bud differentiation and flowering time in L. radiata. These results can preliminarily explain the molecular mechanism of floral bud differentiation and provide new candidate genes for the flowering regulation of Lycoris.

Harnessing phenological traits of wild ancestor Chenopodium hircinum to improve climate adaptation of quinoa

Context Cultivation of quinoa (Chenopodium quinoa Willd.) is rapidly expanding worldwide. Characterisation of populations of Chenopodium hircinum Schard., its wild ancestor, which thrives in some of the hottest environments in South America, may provide adaptations to new environments. Aims This study evaluated the developmental patterns of populations of C. hircinum collected from a range of agroecological environments in Argentina, in order to quantify variability among sites of origin and to explore the association between climatic data from environments of provenance and variation in development. Methods Thirty-three populations of C. hircinum from contrasting sites of origin in Argentina were multiplied in a common-garden experiment under non-limiting conditions of water and nutrient availability. Plants were sampled once or twice weekly (according to parameter) for estimation of the duration of developmental phases, leaf number, and dates of initiation of branching on the main stem. Key results Significant variation was detected for all phenological traits, and populations were categorised into six groups based on similarity of patterns of variation. We found positive association of the duration of development phases and the number of leaves on the main-stem with maximum temperature during the growing season, and negative association with altitude of origin, consistent with variation in growing-season duration. Conclusions The finding that late-flowering populations are associated with warmest climates reveals that longer vegetative growth is an adaptive strategy to cope with heat stress in Chenopodium spp. Implications Time to flowering should be considered in attempts to improve quinoa performance under heat-stress conditions. Further work is needed to understand the genetic basis controlling this response in wild populations of C. hircinum.

Is increased male flower production a strategy for avoidance of predispersal seed predation in andromonoecious plants?

Floral gender in angiosperms often varies within and among populations. We conducted a field survey to test how predispersal seed predation affects sex allocation in an andromonoecious alpine herb Peucedanum multivittatum. We compared plant size, male and perfect flower production, fruit set, and seed predation rate over three years among nine populations inhabiting diverse snowmelt conditions in alpine meadows. Flowering period of individual populations varied from mid‐July to late August reflecting the snowmelt time. Although perfect flower and fruit productions increased with plant size, size dependency of male flower production was less clear. The number of male flowers was larger in the early‐flowering populations, while the number of perfect flowers increased in the late‐flowering populations. Thus, male‐biased sex allocation was common in the early‐flowering populations. Fruit‐set rates varied among populations and between years, irrespective of flowering period. Fruit‐set success of individual plants increased with perfect flower number, but independent of male flower number. Seed predation by lepidopteran larvae was intense in the early‐flowering populations, whereas predation damage was absent in the late‐flowering populations, reflecting the extent of phenological matching between flowering time of host plants and oviposition period of predator moths. Seed predation rate was independent of male and perfect flower numbers of individual plants. Thus, seed predation is a stochastic event in each population. There was a clear correlation between the proportion of male flowers and the intensity of seed predation among populations. These results suggest that male‐biased sex allocation could be a strategy to reduce seed predation damage but maintain the effort as a pollen donor under intensive seed predation.

Utilization of natural variation in Cephalaria joppensis to diversify wheat forage crop rotation in Israel

The recently domesticated species, Cephalaria joppensis (CJ), emerges as a new alternative forage crop in Israel. It has high biomass potential and nutritional values that are comparable to forage wheat. Still, much of the agronomic information regarding CJ is based mainly on a single variety, cv. Rishon, and the genetic variability of this species has not been evaluated. In the last 3 years, CJ seeds have been collected from more than 200 natural populations in Israel. In this work, we characterized 42 of these populations in a replicated field trial, using cv. Rishon as a control. In addition, near infrared spectroscopy (NIRS) was calibrated to predict nutritional attributes. NIRS was found to be instrumental in producing excellent predictions of ash, crude protein, neutral detergent fiber, in-vitro digestibility and rumen degradability, but it did not predict lignin or nitrate. Large variation was found among the accessions with respect to growth rate, flowering time and yield, with several accessions scoring significantly higher than cv. Rishon. Almost no variation was found in nutritional quality-related traits. Early flowering populations were somewhat less fibers and higher digestibility than late flowering populations. The natural variation in agronomic traits will facilitate the development of new breeding germplasm for CJ in the near future.

New species and subspecies of <i>Babiana</i>, <i>Hesperantha</i>, and <i>Ixia</i> (Iridaceae: Crocoideae) from southern Africa; range extensions and morphological and nomenclatural notes on <i>Babiana</i> and <i>Geissorhiza</i>

Babiana rivulicola from stream banks in the Kamiesberg in Namaqualand and terete-leaved Ixia teretifolia from the Roggeveld, both in Northern Cape, are new species of these two largely winter-rainfall region genera. Late-flowering populations of Hesperantha radiata with crowded spikes of smaller flowers are segregated from the typical form as subsp. caricina. We also document the first record of B. gariepensis from Namibia, correct the authority for B. purpurea Ker Gawl., discuss morphologically aberrant populations of B. tubiflora from Saldanha, provide an expanded description for B. lapeirousiodes based on the second and only precisely localized collection of this rare Namaqualand species, and expand the circumscription of Geissorhiza demissa to accommodate a new record from the Kamiesberg, including revised couplets to the existing key to the species.

Large delay in flowering in continental versus coastal populations of a Mediterranean shrub, Globularia alypum

Globularia alypum is a perennial shrub typical of western Mediterranean thermophilous shrublands. Nine populations of G. alypum located in different localities of Catalonia (NE Spain) were surveyed for flowering phenology. Flower-head buds were present in all the populations in July. Flowering time in the area spans from the late summer-early autumn to the next spring depending on the populations; there are two groups of populations, early and late flowering. Early populations grow mostly in coastal localities and flower from September to November, whereas late flowering populations grow in inland localities and flower from February to April. The flowering order of the populations correlated with minimum temperature of most months except the warmest ones, and correlated with maximum and mean temperatures of the coldest months. Correlations were similar when tested with annual climate. The flowering order also correlated with the thermic interval for most months except the coldest and with the index of continentality. Early populations alone did not present correlations with any variable, whereas late populations alone correlated similarly to all populations together. Flowering order did not correlate with precipitation. Late populations are proposed to be regulated by temperature according to our results whereas early populations could be regulated by timing in precipitation after summer drought, according to published results. We discuss the possibilities of the two flowering patterns, early and late, being due to phenotypic plasticity or to genetic adaptation to local climates. We also discuss the consequences at the plant and ecosystem level of climate warming causing shifts from late to early patterns, a possibility that is likely in the warmest of the late populations if flowering is modulated phenotypically.

Can feral weeds evolve from cultivated radish (Raphanus sativus, Brassicaceae)?

Cultivated plants that cannot survive on their own often have maladaptive domestication traits. Unharvested crop seeds may generate feral populations, at times causing serious weed problems, but little is known about the evolution of ferality. We explored the potential for cultivated radish, Raphanus sativus, to become feral, given that closely related taxa (e.g., R. raphanistrum and crop-wild hybrids) are well-documented weeds. First, we measured the population growth of five experimental, cultivated, self-seeding radish populations in Michigan, USA, for three generations. Three late-flowering populations went extinct, and two others apparently hybridized with local R. raphanistrum. A common garden experiment showed that the two surviving populations had earlier flowering, smaller root diameters, and greater individual fecundity than did nonhybridized populations. We also used artificial selection to measure the evolutionary potential for earlier flowering. After two generations of strong selection, two of three lineages flowered earlier and produced more seeds than control lineages, but insufficient genetic variation prevented dramatic evolution of crop phenotypes. In summary, it seems unlikely that radishes could spontaneously become feral in our study area without gene flow from R. raphanistrum. Applying these approaches to other cultivated species may provide a better understanding of mechanisms promoting the evolution of feral weeds.

Natural Variation in Arabidopsis lyrata Vernalization Requirement Conferred by a FRIGIDA Indel Polymorphism

Species share homologous genes to a large extent, but it is not yet known to what degree the same loci have been targets for natural selection in different species. Natural variation in flowering time is determined to a large degree by 2 genes, FLOWERING LOCUS C and FRIGIDA, in Arabidopsis thaliana. Here, we examine whether FRIGIDA has a role in differences in flowering time between and within natural populations of Arabidopsis lyrata, a close outcrossing perennial relative of A. thaliana. We found 2 FRIGIDA sequence variants producing potentially functional proteins but with a length difference of 14 amino acids. These variants conferred a 15-day difference in flowering time in an association experiment in 2 Scandinavian populations. The difference in flowering time between alleles was confirmed with transformation to A. thaliana. Because the north European late-flowering populations harbor both late- and early sequence variants at intermediate frequencies and the late-flowering variant is most frequent in the southern early flowering European population, other genetic factors must be responsible for the flowering time differences between the populations. The length polymorphism occurs at high frequencies also in several North American populations. The occurrence of functional variants at intermediate frequencies in several populations suggests that the variation may be maintained by balancing selection. This is in contrast to A. thaliana, where independent loss-of-function mutations at the FRIGIDA gene are responsible for differences between populations and local adaptation.

Little genetic differentiation across Europe between early-flowering and late-flowering populations of the rapidly declining orchid Neotinea ustulata

Amplified fragment length polymorphism (AFLP) was used to characterize genetic diversity of the endangered Burnt Orchid, Neotinea (formerly Orchis) ustulata. Fingerprinting of Estonian and British populations revealed surprisingly little genetic differentiation between populations but larger amounts of diversity within populations, especially in Britain. The resulting mean Fst value of 0.51 is unusually high for an orchid species. Much of the variation follows a west–east cline across Europe, whereas the much-discussed early- and late-flowering taxa of N. ustulata are considered insufficiently distinct to be viewed as separate subspecies. The later flowering N. ustulata var. aestivalis probably evolved independently on two or three occasions, each time diverging from the earlier flowering nominate race. The identity of the genes underpinning phenology in the species, and the potential selective advantages of phenological divergence, merit further study. Overall genetic diversity within populations is sufficiently high to render impoverishment an unlikely cause of their recent, precipitous decline. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87, 13–25.

Morphometric and genetic divergence among populations ofNeotinea ustulata (Orchidaceae) with different flowering phenologies

The terrestrial orchid speciesNeotinea ustulata has recently been split into two subspecies, differing remarkably in their flowering time, but only slightly in morphological characteristics, which makes their taxonomic status uncertain. We have analyzed morphometric and genetic differences between the early- and late-flowering populations in Central Europe. Our results on morphology are ambiguous. Indirect gradient analysis has not shown a distinct separation of early- and late-flowering individuals in the ordination space. However, according to MANOVA, populations of early- and late-flowering plants can be distinguished by plant height, leaf length, numbers of basal (rosette) and stem leaves and even better by certain ratios of these numbers. All genetic analyses, on the other hand, are definite and consistently distinguish two groups. Random amplified polymorphic DNA (RAPD) markers have shown that the early- and late-flowering populations differ significantly from one another. Principal coordinate analysis (PCoA) based on presence/absence matrix of RAPD bands separated the two groups, implying that the difference in flowering phenology could form an effective barrier to gene exchange. Partitioning of genetic diversity in analysis of molecular variance (AMOVA) has shown that the genetic divergence between the two groups, early- and late-flowering populations, is somewhat greater (33%) than the genetic variability among populations within particular group (23%). Using the Mantel test, we found that genetic differentiation coefficients between populations closely correspond to their geographic distribution. After elimination of the effect of sample origin from the model, direct gradient analysis (RDA) has shown that the early- and late-flowering groups differ significantly in their RAPD spectra. To conclude, our results indicate the presence of two genetically and phenologically distinct taxa, but the weak morphological differentiation supports the taxonomic rank of variety rather than subspecies.

Highly variable flowering time in Orchis ustulata (Orchidaceae): consequences for population dynamics

Two subspecies of Orchis ustulata differ considerably in their flowering time, but in only a few morphometric parameters; this makes the rank of these taxa problematic. The flowering time was not affected by replanting individuals to another habitat, including to the habitat of the other subspecies. In this study, populations from both subspecies were studied, measuring 464 marked genets during 5–6 years. Populations with different flowering times exhibited notable differences in their local distribution areas and the mean height of specimens. In the late‐flowering populations the proportion of dormant plants is higher and the proportion of vegetatively propagated shoots lower than in the early‐flowering ones. Going to (or staying) dormant is the biggest possibility in all stage groups. Flowering is more likely to be followed by dormancy than vegetative stage, but setting fruit does not affect the possibility. Vegetative propagation may play an important role in keeping the populations viable. Vegetative growth is more pronounced on stony soils.

Tolerance of Gentianella campestris in relation to damage intensity: an interplay between apical dominance and herbivory

Meristem allocation models suggest that the patterns of compensatory regrowth responses following grazing vary, depending on (i) the number of latent meristems that escape from being damaged, and (ii) the activation sensitivity of the meristems in relation to the degree of damage. We examined the shape of compensatory responses in two late-flowering populations (59°20′N and 65°45′N) of the field gentian. Plants of equal initial sizes were randomly assigned to four treatment groups with 0, 10, 50 and 75% removal of the main stalk. The plants were clipped before flowering, and their performance was studied at the end of the growing season. The northern population showed a linear decrease in shoot biomass and fecundity with increasing biomass removal, while the response in the southern population was quadratic with maximum performance at the damage level of 50% clipping. This nonlinear shape depended upon the activation sensitivity of dormant meristems in relation to their position along the main stem. The highest plant performance was achieved by inflicting intermediate damage which induced regrowth from basally located meristems. In contrast, the topmost branches took over the dominance role of the main stem after minor apical damage (10% clipping). Consequently, the breakage of apical dominance is a necessary precondition of vigorous regrowth in this species. However, compensation in the field gentian is unlikely to be a mere incidental by-product of apical dominance. The ability to regrow from basally located meristems that escape from being damaged by grazing may well be a sign of adaptation to moderate levels of shoot damage.

Genetic and Floral Divergence among Sympatric Populations of Gymnadenia conopsea s.l. (Orchideaceae) with Different Flowering Phenology.

Gymnadenia conopsea s.l. is a common orchid in central Europe, where early- and late-flowering populations can be distinguished. The early-flowering form is recognized as subspecies conopsea and the late-flowering form as subspecies densiflora. The two subspecies can occur in sympatry, but their flowering periods are separated. We investigated whether early- and late-flowering subspecies are genetically differentiated, whether they diverged once or repeatedly, and we tried to identify potential evolutionary forces involved in the divergence of the two subspecies. We used genetic markers to estimate genetic divergence within and among populations of early- and late-flowering G. conopsea, and to reconstruct their evolutionary history. In addition, we assessed morphological variation between subspecies. Allozyme variation indicated that subspecies conopsea was significantly more variable than ssp. densiflora and that gene flow among populations of ssp. conopsea was higher than among populations of ssp. densiflora. Gene flow between subspecies was low, indicating that the difference in flowering phenology represented an effective barrier to gene flow. A neighbor-joining tree based on allozyme frequencies indicated that early- and late- flowering populations did not diverge repeatedly in sympatry. Levels of cpDNA variation were generally low, even between G. conopsea s.l. and Gymnadenia odoratissima, chosen as an outgroup. Four cpDNA haplotypes were found, which differed only in the number of microsatellite repeats. Their distribution among subspecies of G. conopsea s.l. and G. odoratissima indicates that microsatellite haplotypes have evolved repeatedly, and their occurrence in different taxa thus represents a homoplasy. Floral characters were variable within and among populations and subspecies but did not consistently separate early- from late-flowering populations. A weak separation between subspecies was found in vegetative characters that presumably reflected habitat and competitive differences experienced by early- and late-flowering populations.

Variability in habit, flowering and seed production within the Kangaroo Valley cultivar of Lolium perenne when grown in a range of environments

Diversity of flowering and seed production characteristics were measured within and between seven populations of Kangaroo Valley perennial ryegrass. The hypothesis that these characteristics would not change when grown in diverse environments was tested. The average time from seedling emergence to spike emergence was similar for each of three biotypes that represented early, intermediate and late flowering populations. This occurred in spite of variation in this character of 54 to 72 days between plants within each of these biotypes. When data from all seven populations were combined, they formed a normal distribution with respect to time of flowering. Distinctive components of seed production were found for the different populations. Early flowering populations had higher seed yields and more but shorter spikes than late flowering populations. Wide variation in habit occurred within populations but there was a high correlation between erect habit and early flowering. Spikes of erect plants emerged 27 days earlier than prostrate plants. The seed formation characteristics of an early flowering population were unaltered when grown for three generations in diverse environments. The spike emergence and anthesis times for each of the seven populations were not changed when grown in new environments. We conclude that seed production of Kangaroo Valley ryegrass in environments similar to those described in these experiments will not significantly alter the population characteristic. For changes in the character of the population to occur, the new environment would have to be so severe to (a) restrict growth and select out a proportion of the progeny or (b) cause a change in the time of seed harvest.