SPATIAL AND TEMPORAL VARIATION IN CHASMOGAMY AND CLEISTOGAMY IN OXALIS MONTANA (OXALIDACEAE)

Chasmogamous (CH) and cleistogamous (CL) flower production was investigated in natural populations of the perennial herb Oxalis montana in southern Quebec, Canada. Every 10–12 days during two flowering seasons, we recorded the reproductive output of approximately 2,000 plants distributed among five forest sites. The percentage of plants flowering, proportion of flowering plants producing CH and CL flowers, CH and CL flower number per flowering plant, and the proportion of flowers that are CL differed significantly among sites and corresponded to site differences in forest type and habitat quality. Unlike patterns in most CL species, however, reproduction by cleistogamy increased in sites with habitat conditions favorable to plant growth and sexual reproduction, and decreased in less productive sites. Chasmogamous and CL flower production increased with increasing plant size but plant size explained a significantly greater proportion of the variation in CL flower numbers. The percentage of flowering plants producing CH flowers decreased between years while the proportion of CL flowers increased at all sites during the second flowering season. The somewhat unusual aspects of reproduction in Oxalis may stem from its perennial habit which allows use of stored resources in adjusting the balance of cleistogamy versus chasmogamy in different environmental regimes.     

Reproductive ecology of Viola mirabilis var. subglabra representing intermediate flowering characteristics between stemless and stemmed Viola

Viola (Violaceae) is one of the largest genera in angiosperms. This genus is essentially classified into stemless and stemmed groups based on growth morphology. However, Viola mirabilis var. subglabra is an exception in having intermediate flowering characteristics; cleistogamous (CL) flowers are formed in the axils of stem leaves, whereas chasmogamous (CH) flowers arise from basal rosettes (radical CH (CH(r)) flowers) and also in the axils of the stem (axially CH (CH(a)) flowers). To understand why the pattern of flower production varies in this Viola species, flower production was investigated in 10 Japanese populations from Hokkaido to the western part of Honshu in 2014 and 2015. Furthermore, flower characteristics were also compared between CH(r) and CH(a) flowers in Hokkaido. In this species, the production of CH flowers varied among individuals, and they were categorized into three groups, individuals that produced (i) only CH(r) flowers, (ii) only CH(a) flowers and (iii) both CH flowers. The frequency of these groups differed among populations, but some individuals changed the category between 2014 and 2015. Thus, the production of CH(r) and CH(a) flowers plastically changes depending on individual conditions and/or environmental factors. On the other hand, CH(r) and CH(a) flowers differed in flower size and flowering phenology. These results suggest that two types of CH flowers may play different roles in reproduction in each population, but fruit sets and seed sets did not differ between two types of CH flowers.     

Why is there variation in mean seed size among plants within single populations? test of the fertilization efficiency hypothesis

We tested the fertilization efficiency hypothesis, which attempts to explain mean seed size variation among plants within single populations, by comparing the patterns of seed size variation between chasmogamous (CH) flowers and cleistogamous (CL) flowers in Impatiens noli-tangere and Viola grypoceras, respectively. The fertilization efficiency hypothesis predicts that larger plants produce larger seeds if the number of pollen grains captured by a plant increases with increased allocation of resources to its attractive structures (e.g., corolla and nectar), but with diminishing gains. Thus, seed size should depend on plant size in seeds from CH flowers because of the diminishing gains of capturing pollen in these flowers, whereas seed size should not depend on plant size in seeds from CL flowers because CL flowers need not capture outcross pollen. We found significant positive correlations between mean seed size per plant and plant size for seeds from CH flowers in both species. However, there was no significant positive correlation between these two factors for seeds from CL flowers of both species. The results of the present investigations were thus consistent with the fertilization efficiency hypothesis.     

THE BALANCE BETWEEN CHASMOGAMY AND CLEISTOGAMY IN COLLOMIA GRANDIFLORA (POLEMONIACEAE)

The extent of cleistogamy (CL) and chasmogamy (CH) was investigated in natural populations of Collomia grandiflora Dougl. ex Lindl. The ubiquitous occurrence of CL flowers was documented in 18 populations from throughout the geographic range of the species. The interrelationship of environmental perturbation, vegetative weight, and the balance between CH and CL flowers was investigated by means of a factorial experiment which varied plant density, incident light and soil mix. An ANOVA revealed that total plant weight increased significantly under conditions of low density, low light and low sand content, with density accounting for most of the variance in plant weight. Interactions involving density were not significant. The number of CH and CL flowers and estimated percent outcrossing increased significantly with increased plant weight, low density, low light, and low sand content, although an analysis of covariance showed that plant weight and density accounted for most of the variance in these traits. Among large, multibranched plants, the proportion of CL flowers increased in a basipetal pattern, whereas most of the CH flowers were restricted to the terminal and upper inflorescences. The smallest plants produced only a single terminal inflorescence comprising entirely CL flowers. The proportion of reproductive effort allocated to outcrossing was dependent upon plant size as estimated from total weight, whereas the proportion allocated to ***selfing was relatively independent. Studies of outcrossed vs. inbred progenies of five populations revealed little evidence for immediate inbreeding depression with respect to plant weight and the number of CH or CL flowers, but indicated that significant quantitative genetic differences exist for these traits among populations of diverse ecological or geographical origin.     

EFFECTS OF WATER STRESS,ABSCISIC ACID,AND GIBBERELLIC ACID ON FLOWER PRODUCTION AND DIFFERENTIATION IN THE CLEISTOGAMOUS SPECIES COLLOMIA GRANDIFLORA DOUGL. EX LINDL. (POLEMONIACEAE)

The effects of water stress, abscisic acid (ABA), and gibberellic acid (GA₃) on flower production and differentiation by Collomia grandiflora were investigated. An untreated plant typically produced both small, closed cleistogamous (CL) and large, open chasmogamous (CH) flowers. The larger corolla of CH flowers was due to a greater cell number and size. When plants were water-stressed or sprayed with ABA, both the percentage of CH flowers and the total number of flowers were reduced significantly. The corolla dimensions and epidermal cell numbers and sizes of CL flowers produced by water-stressed and ABA-sprayed plants did not differ from those of CL flowers produced by control plants. Application of GA₃ to both well-watered and water-stressed plants significantly increased the percentage of CH flowers formed compared to well-watered controls. In the absence of GA₃, water-stressed plants produced almost entirely CL flowers. GA₃-sprayed plants produced CH flowers whose corolla dimensions were intermediate between those of CL and CH flowers formed by control plants. Epidermal cells of these intermediate corollas were reduced only in number and not in size when compared to control CH flowers. Endogenous levels of ABA and gibberellins may control the type of flower produced by C. grandiflora and may mediate some of the observable effects of water stress on flowering.     

The Life History of Euryale ferox Salisb. in Southwestern Japan with Special Reference to Reproductive Ecology

Abstract The life history of Euryale ferox Salisb, observed in Akashi, Southwestern Japan, is described. Cleistogamous (CL) flowers appear from early July and precocious pollination occurs. Chasmogamous (CH) flowers are seen during a limited period between mid-August and mid-September. The abundance of CH flowers varies among populations and/or from year to year. Few insects visit CH flowers and self-pollination occurs prior to anthesis. The number of seeds produced by CL flowers significantly exceeds that of CH flowers. Considering the shorter period of CH flower production and the lesser quantity of seed produced, the significance of CL flowers in the maintenance of E. ferox populations must be considered greater than that of CH flowers.     

Sex expression and breeding strategy in <Emphasis Type="Italic">Commelina benghalensis</Emphasis> L.

This paper describes the results of a series of experiments conducted to unravel the patterns of sex expression and reproductive output in a fascinating species with high variation in sexuality. Commelina benghalensis L., an andromonoecious rainy season weed, bears male and bisexual flowers in axillary spathes of all the plants investigated. Bisexual flowers are of two types; chasmogamous (CH) and cleistogamous (CL). The former are borne on subaerial and the latter on subterranean shoots, in addition to those on aerial spathes. Three populations of the species, designated JU1, JU2 and JU3, were scanned for three consecutive years from 1996 to 1998, and the number and distribution of male, CH and CL flowers per plant were found to vary. The mere number of CH/CL flowers per plant is by itself not an accurate measure of mixed mating. It is necessary to confirm that CH flowers actually outcross and, if they do so, to what extent. Comparison of the pollen/ovule (P/O) ratio and percentage pollen germination on the stigmas of the CH and CL flowers have been used as indices of the pollination system. Confirmation of this was sought from the fruit and seed sets obtained after manual pollination of emasculated flowers with self- and cross-pollen. Results so obtained were compared with those of natural pollination. In the majority of CH flowers, the male and female reproductive phases (i.e. anther dehiscence and stigma receptivity) overlap, providing for self-pollination. However, two exceptions to this general behaviour were found in some plants of all the three populations. In some CH flowers, the female phase matures prior to anther dehiscence while in others, the anthers are sterile. Such plants, designated as variants 1 and 2, respectively, facilitate cross-pollination. While the CL flowers contribute to the production of selfed progeny, the variants of CH ones permit formation of outcrossed progeny, indicating a mixed mating strategy in C. benghalensis.     

Unpredictability of nectar nicotine promotes outcrossing by hummingbirds in Nicotiana attenuata

Many plants use sophisticated strategies to maximize their reproductive success via outcrossing. Nicotiana attenuata flowers produce nectar with nicotine at concentrations that are repellent to hummingbirds, increasing the number of flowers visited per plant. In choice tests using native hummingbirds, we show that these important pollinators learn to tolerate high‐nicotine nectar but prefer low‐nicotine nectar, and show no signs of nicotine addiction. Nectar nicotine concentrations, unlike those of other vegetative tissues, are unpredictably variable among flowers, not only among populations, but also within populations, and even among flowers within an inflorescence. To evaluate whether variations in nectar nicotine concentrations increase outcrossing, polymorphic microsatellite markers, optimized to evaluate paternity in native N. attenuata populations, were used to compare outcrossing in plants silenced for expression of a biosynthetic gene for nicotine production (Napmt1/2) and in control empty vector plants, which were antherectomized and transplanted into native populations. When only exposed to hummingbird pollinators, seeds produced by flowers with nicotine in their nectar had a greater number of genetically different sires, compared to seeds from nicotine‐free flowers. As the variation in nectar nicotine levels among flowers in an inflorescence decreased in N. attenuata plants silenced in various combinations of three Dicer‐like (DCL) proteins, small RNAs are probably involved in the unpredictable variation in nectar nicotine levels within a plant.     

Spatio‐temporal patterns in pollination of deceptive Aristolochia rotunda L. (Aristolochiaceae)

• Pollination success of highly specialised flowers is susceptible to fluctuations of the pollinator fauna. Mediterranean Aristolochia rotunda has deceptive trap flowers exhibiting a highly specialised pollination system. The sole pollinators are kleptoparasitic flies in search of food. This study investigates these pollinators on a spatio‐temporal scale and the impact of weather conditions on their availability. Two potential strategies of the plants to cope with pollinator limitation, i.e. autonomous selfing and an increased floral life span, were tested.
• A total of 6156 flowers were investigated for entrapped pollinators in 10 Croatian populations. Availability of the main pollinator was correlated to meteorological data. Artificial pollination experiments were conducted and the floral life span was recorded in two populations according to pollinator availability.
• Trachysiphonella ruficeps (Chloropidae) was identified as dominant pollinator, along with less abundant species of Chloropidae, Ceratopogonidae and Milichiidae. Pollinator compositions varied among populations. Weather conditions 15–30 days before pollination had a significant effect on availability of the main pollinator. Flowers were not autonomously selfing, and the floral life span exhibited considerable plasticity depending on pollinator availability.
• A. rotunda flowers rely on insect pollen vectors. Plants are specialised on a guild of kleptoparasitic flies, rather than on a single species. Pollinator variability may result in differing selection pressures among populations. The availability/abundance of pollinators depends on weather conditions during their larval development. Flowers show a prolonged trapping flower stage that likely increases outcrossing success during periods of pollinator limitation.

     

Patterns of flower morphology and structural changes during interconversion between chasmogamous and cleistogamous flowers in Viola philippica北大核心CSCD

Aims: Viola philippica is a species with a typical chasmogamous-cleistogamous (CH-CL) mixed breeding system. It provides a flower model system to investigate floral organs development under different photoperiods. Morphological changes of intermediate cleistogamous (inCL) flowers have been observed, the trends in variation of changes from CH flowers to CL flowers or from CL flowers to CH flowers have been analyzed, the localized effects of poorly developed stamens and petals in CL and inCL flowers have been identified. This research provided morphology and structural changes with implication for the evolutionary significance of the dimorphic flower formation for further study in dimorphic flower development. Methods: We used methods of anatomy and structural analysis to observe the morphological structures of flowers under different photoperiods. Important findings: Photoperiod played an important role in the development of CH and CL flowers in V. philippica. Under short-day light and intermediate-day light, both CH and inCL flowers developed simultaneously. Most of the floral buds were CH flowers under a photoperiod of short-day light, but most of the floral buds were inCL flowers under mid-day light. Complete CL flowers formed under long-day lights. However, there were a series of transitional types in the number and morphology of stamens and petals among inCL flowers, including five stamens with three petals related to CH flowers and two stamens with one petal related to CL flowers. The former type was dominant under short-day light conditions, and the latter type was dominant under mid-day light. Further more, there were localized effects in stamen and petal development for CL and inCL flowers. The development of ventral lower petal (corresponding to the lower petal with spur of CH flower) and the adjacent two stamens in inCL flowers were best, and the back petal was similar to that of CL flowers, an organ primordium structure. The adjacent stamens with the back petals tended to be poorly developed. In extreme cases, these stamens in inCL flowers had no pollen sac, only a membranous appendage or even a primordium structure. When the plants with CL or CH flowers were placed under short-day light or long-day light, the newly induced flowers all showed a series of inCL flower types, finally the CL flowers transformed into CH flowers, and the CH flowers transformed into CL flowers. This result indicates the gradual effects of different photoperiods on dimorphic flowers development of V. philippica. A long photoperiod could inhibit the development of partial stamens and petals, and a short photoperiod could prevent the suppression of long-day light and promote the development of stamens and petals.     

Life history trade‐offs and evidence for hierarchical resource allocation in two monocarpic perennials

The evolution of floral display is thought to be constrained by trade‐offs between the size and number of flowers; however, empirical evidence for the trade‐off is inconsistent. We examined evidence for trade‐offs and hierarchical allocation of resources within and between two populations each of the monocarpic perennials, Cardiocrinum cordatum and C. giganteum. Within all populations, flower size–number trade‐offs were evident after accounting for variation in plant size. In addition, variation in flower size explained much variation in flower‐level allocation to attraction, and female and male function, a pattern consistent with hierarchical allocation. However, between population differences in flower size (C. cordatum) and number (C. giganteum) were not consistent with size–number trade‐offs or hierarchical allocation. The population‐level difference in C. cordatum likely reflects the combined influence of a time lag between initiation and maturation of flowers, and higher light levels in one population. Thus, our study highlights one mechanism that may account for the apparent independence of flower size and number in many studies. A prediction of sex allocation theory was also supported. In C. giganteum: plants from one population invested more mass in pistils and less in stamens than did plants from the other population. Detection of floral trade‐offs in Cardiocrinum may be facilitated by monocarpic reproduction, production of a single inflorescence and ease of measuring plant size.     

ENHANCEMENT OF INBREEDING DEPRESSION BY DOMINANCE AND SUPPRESSION IN IMPATIENS CAPENSIS

We investigated the effect of intraspecific competition on the magnitude of inbreeding depression in Impatiens capensis by planting seeds from chasmogamous (CH) and cleistogamous (CL) flowers in three experimental greenhouse treatments: in individual pots, in flats in dense pure stands according to seed type, and in flats with the two seed types intermixed in a checkerboard array. The size distributions of plants grown in flats were significantly more hierarchical than those of plants grown individually, indicating that larger plants competitively suppressed smaller plants in the high-density treatments. The magnitude of inbreeding depression at high density depended upon the planting arrangement of CL and CH seeds. CH advantage was greatest when CH and CL seedlings were grown in competition with one another, suggesting that fitness differences between outcrossed and inbred individuals were intensified by dominance and suppression. For plants grown individually, the effects of maternal parent, seed weight, and emergence date on seedling size disappeared with plant age, whereas at high density these effects remained at the final harvest. Thus, plant density may influence patterns of natural selection both on mating system and on juvenile traits in natural Impatiens populations.     

Phenotypic selection in a natural population of Impatiens pallida Nutt. (Balsaminaceae)

In the annual plant Impatiens pallida, individuals exhibit a floral heteromorphism consisting of autogamously selfing, cleistogamous (CL) flowers and partially outcrossing, chasmogamous (CH) flowers. As part of an investigation into natural selection and mating system evolution in I. pallida, we measured the magnitude and direction of phenotypic selection on nine life history characters (two traits measured on three dates, one measured on two dates and one measured once). Three of these characters were positively correlated with the ratio of CH/CL flowers produced per plant, which is an important determinant of the mating system. Values for the nine characters and three different measures of fitness (viability, fecundity, lifetime) were estimated for 500 plants in five locations over a single growing season. Based on lifetime fitness, linear selection differentials were significant for all nine characters, indicating a selective advantage to tall, leafy, highly branched plants that flowered early. However, only two of these characters had a direct effect on fitness. Selection was significant on all nine characters when based on fecundity as well as lifetime fitness; however, only three of five characters examined had significant selection based on viability fitness. For all fitness components, the frequency of significant linear and nonlinear selection coefficients was comparable (23% vs 17% of all cases, respectively), but nonlinear coefficients were generally larger. Finally, the magnitude and direction of direct linear selection was heterogeneous among locations, for all characters and all fitness components. Collectively these results suggest that selection is strong, favouring large size, high allocation to reproduction and high CH/CL flower ratios. However, any directional evolutionary changes in vegetative or reproductive characters may be constrained by strong non-linear and correlational selection.     

Effects of cleistogamy and pollen source on seed production and offspring performance in three endangered violets

In rare plants that often occur in small or isolated populations the probability of selfing between close relatives is increased as a consequence of demographic stochasticity. The mode of pollination (selfing, outcrossing) may have considerable effects on seed traits and offspring performance and hence potential viability. Since current efforts aiming at the restoration of floodplain grasslands through the transfer of plant material from species-rich source stands may lead to the establishment of initially small populations consisting of founders from different populations, the present paper experimentally investigated the effects of pollen source and floral types (i.e. chasmogamous (CH) and cleistogamous (CL) flowers) on seed traits and offspring performance in three highly endangered violet species (Viola elatior, V. pumila, V. stagnina) of these grasslands. We estimated inbreeding depression and tested the performance of selfed and outcrossed offspring in two microbial environments, i.e. in soil inoculated with (i) non-sterile substrate from the same species (‘home’-conditions) and (ii) sterilised substrate.Plants produced more CL capsules than CH flowers. Pollinator exclusion had only small effects on CH seed production. CL seeds had a significantly lower mass per seed than CH seeds. This may be related to constraints in allocation or environmental conditions. Seedling growth was reduced in plants grown under ‘home’-conditions as compared to control soils. Under ‘home’-conditions, relative fitness of selfed seedlings of V. stagnina was significantly higher than that of crossed progeny. Our results suggest that high genetic differentiation among populations as a consequence of isolation may result in outbreeding depression, e.g., through biochemical or physiological incompatibilities between genes or the breaking of coadapted gene complexes. In V. stagnina, offspring fitness differed considerably between environments, but in general we found no indications for inbreeding depression in these rare species.     

CHASMOGAMY/CLEISTOGAMY IN TRIODANIS PERFOLIATA (CAMPANULACEAE): SOME CH/CL COMPARISONS IN FITNESS PARAMETERS

The chasmogamous/cleistogamous (CH/CL) breeding system has evolved independently many times, but its ecological role has been studied in few species. In Triodanis perfoliata, we studied the natural phenology of flowering and seed set, how successful CH flowers were at setting seed, whether seed from CH and CL flowers differed in germination behavior, and whether plants derived from these two seed types differed in vigor. Most notably, we found that the later the CH flowers were produced, the fewer seed they successfully matured. In general the two seed types did not show differences in germination behavior, but for some maternal parents the two seed types differed. For these, the CL seed germinated less readily. Finally, there was some indication that plants from CL seed were not as vigorous as those from CH seed.     

Patterns of flower morphology and structural changes during interconversion between chasmogamous and cleistogamous flowers in Viola philippica

Aims Viola philippica is a species with a typical chasmogamous-cleistogamous (CH-CL) mixed breeding system. It provides a flower model system to investigate floral organs development under different photoperiods. Morphological changes of intermediate cleistogamous (inCL) flowers have been observed, the trends in variation of changes from CH flowers to CL flowers or from CL flowers to CH flowers have been analyzed, the localized effects of poorly developed stamens and petals in CL and inCL flowers have been identified. This research provided morphology and structural changes with implication for the evolutionary significance of the dimorphic flower formation for further study in dimorphic flower development.Methods We used methods of anatomy and structural analysis to observe the morphological structures of flowers under different photoperiods.Important findings Photoperiod played an important role in the development of CH and CL flowers in V. philippica. Under short-day light and intermediate-day light, both CH and inCL flowers developed simultaneously. Most of the floral buds were CH flowers under a photoperiod of short-day light, but most of the floral buds were inCL flowers under mid-day light. Complete CL flowers formed under long-day lights. However, there were a series of transitional types in the number and morphology of stamens and petals among inCL flowers, including five stamens with three petals related to CH flowers and two stamens with one petal related to CL flowers. The former type was dominant under short-day light conditions, and the latter type was dominant under mid-day light. Further more, there were localized effects in stamen and petal development for CL and inCL flowers. The development of ventral lower petal (corresponding to the lower petal with spur of CH flower) and the adjacent two stamens in inCL flowers were best, and the back petal was similar to that of CL flowers, an organ primordium structure. The adjacent stamens with the back petals tended to be poorly developed. In extreme cases, these stamens in inCL flowers had no pollen sac, only a membranous appendage or even a primordium structure. When the plants with CL or CH flowers were placed under short-day light or long-day light, the newly induced flowers all showed a series of inCL flower types, finally the CL flowers transformed into CH flowers, and the CH flowers transformed into CL flowers. This result indicates the gradual effects of different photoperiods on dimorphic flowers development of V. philippica. A long photoperiod could inhibit the development of partial stamens and petals, and a short photoperiod could prevent the suppression of long-day light and promote the development of stamens and petals.     

CLEISTOGAMY IN MICROLAENA POLYNODA (GRAMINEAE): AN EXAMINATION OF SOME MODEL PREDICTIONS

A population of the cleistogamous grass, Microlaena polynoda, was investigated to determine some of the factors responsible for adjusting the balance between reproduction by chasmogamous (CH) and cleistogamous (CL) flowers, the tissue costs associated with the two reproductive modes, the fate of progeny produced by each mode, and the genetic diversity of the progeny. Cleistogamous flower production begins earlier in the season than CH flower production. There is a distinct threshold of low light intensity below which mostly CL flowers are produced. Paternal expenditure per plant is an order of magnitude larger for the CH than the CL component. The opposite relation holds for maternal expenditure. Increased maternal expenditure in the CL component may be due to greater fertilization success and retrieval of paternal costs. Cleistogamous seeds are dispersed later than CH seeds. Following dispersal, the spikelet encloses the CL seed but not the CH seed, and is responsible for inducing dormancy. The ratio of seedlings arising from CH seeds to that from CL seeds in a natural habitat is significantly lower than the ratio of estimated numbers of CH to CL seeds produced. There were no detectable polymorphisms among ten presumptive enzyme loci assayed. Many of the features associated with CH and CL reproduction in M. polynoda are in accord with the theoretical requirements for the evolution of closed flower self-pollination and the maintenance of two distinct methods of reproduction.     

QUANTITATIVE VARIATION OF PROGENY FROM CHASMOGAMOUS AND CLEISTOGAMOUS FLOWERS IN THE GRASS DANTHONIA SPICATA

Progeny from chasmogamous (CH) and cleistogamous (CL) flowers of the grass Danthonia spicata were raised in their native habitat and in the greenhouse in order to determine how genetic variation was distributed among families and between CH and CL progeny within families. Twelve quantitative characters were measured on clones from individuals known to have arisen from either CH or CL flowers on a particular plant. Significant genetic variation existed for all characters measured. Most genetic variation was between families and two morphologically similar groups of families were identified. Relatively little genetic variation was found within families (approximately 5% of the total phenotypic variance). In field-raised plants, variance component analysis suggested that CL progeny were genetically more similar to each other than were CH progeny from the same plant. Levene's test of the average deviation of CH and CL progenies from their group means was nonsignificant but suggested there was a trend (0.05 < P < 0.10) for CH progeny to be more variable than CL progeny in the field but less variable in the greenhouse. The amount and distribution of genetic variation in the study population indicates that selective differentials would be larger among families than within families.     

Effects of flower dimorphism and light environment on arbuscular mycorrhizal colonisation in a cleistogamous herb

Although it is known that floral dimorphism contributes to the maintenance of mixed breeding systems, the consequences of producing progeny of a contrasting genetic background and seeds with differential resource allocation has been practically ignored regarding establishment of belowground organisms–plant interactions. This article evaluates the combined effect of floral dimorphism with cross type and light environment on interactions between Ruellia nudiflora and arbuscular mycorrhizal fungi (AMF). R. nudiflora produces cleistogamous (CL) flowers that exhibit obligate self‐pollination and chasmogamous (CH) flowers with facultative self‐ (CHs) or cross‐ (CHc) pollination. We evaluated the establishment of the plant–AMF interaction in progeny derived from each floral type, under two light conditions (shaded versus open). We established different scenarios depending on the existence of inbreeding depression (ID) and whether the differential resource allocation (DRA) to CH and CL flowers affected the R. nudiflora–AMF interaction. We predicted that under shaded light conditions there might be an intensification of ID, having a negative effect on AMF colonisation. The percentages of hyphae and vesicles in the harvested roots was significantly higher in the shaded plants (F ≥ 4.11, P < 0.05), while progeny of CHc and CHs presented a higher percentage of hyphae and vesicle colonisation compared to CL progeny (F = 15.26, P < 0.01). The results show that DRA to CH flowers and light availability both determines the establishment of R. nudiflora–AMF interaction. The results also suggest that even under stressful light conditions, endogamy does not affect this interaction, which may explain the success of R. nudiflora as an invasive species.     

Seed germination and seedling allogamy in Rosmarinus officinalis: the costs of inbreeding

• Self‐pollination by geitonogamy is likely in self‐compatible plants that simultaneously expose a large number of flowers to pollinators. However, progeny of these plants is often highly allogamous. Although mechanisms to increase cross‐pollination have been identified and studied, their relative importance has rarely been addressed simultaneously in plant populations.
• We used Rosmarinus officinalis to explore factors that influence the probability of self‐fertilisation due to geitonogamy or that purge its consequences, focusing on their effects on seed germination and allogamy rate. We experimentally tested the effect of geitonogamy on the proportion of filled seeds and how it influences germination rate. During two field seasons, we studied how life history and flowering traits of individuals influence seed germination and allogamy rates of their progeny in wild populations at the extremes of the altitudinal range. The traits considered were plant size, population density, duration of the flowering season, number of open flowers, flowering synchrony among individuals within populations and proportion of male‐sterile flowers.
• We found that most seeds obtained experimentally from self‐pollination were apparently healthy but empty, and that the proportion of filled seeds drove the differences in germination rate between self‐ and cross‐pollination experiments. Plants from wild populations consistently had low germination rate and high rate of allogamy, as determined with microsatellites. Germination rate related positively to the length of the flowering season, flowering synchrony and the ratio of male‐sterile flowers, whereas the rate of allogamous seedlings was positively related only to the ratio of male‐sterile flowers.
• Rosemary plants purge most of the inbreeding caused by its pollination system by aborting the seeds. This study showed that the rates of seed germination and allogamy of the seedlings depend on a complex combination of factors that vary in space and time. Male sterility of flowers, length of the flowering season and flowering synchrony of individuals within populations all favour high rates of cross‐pollination, therefore increasing germination and allogamy rates. Flowering traits appear to be highly plastic and respond to local and seasonal conditions.