The role of a mixed mating system in the reproduction of a Mediterranean subshrub (Fumana hispidula, Cistaceae)

Knowledge about mixed mating systems can improve our understanding of the evolutionary dynamics of reproductive systems. Here we report a study of the pollination system (floral and reproductive biology, pollen limitation and stigmatic pollen load, floral visitors and inbreeding depression) of Fumana hispidula, a Mediterranean subshrub belonging to a species group with a strong selfing tendency. Autonomous self-pollination, hand self- and hand cross-pollination, open pollination and controls yielded fruits (0.28, 0.65, 0.68, 0.38, and 0.67, respectively); however, autonomous self-pollination resulted in the lowest fruit set. All individuals were fully self-compatible but we found great inter-individual variation in autonomous self-pollination ability, which was probably related to a variable expression of herkogamy degree. Inbreeding depression was low (0.040 for fruit set, 0.015 for seed set and ?0.026 for seed mass). The pollen supplementation experiment did not reveal pollen limitation and pollinators were seen visiting the flowers during the observation periods. These results support the idea that F. hispidula has a mixed mating strategy, which represents a successful reproduction mode in their patchy habitats.     

Pollination biology in an endangered rocky mountain toadflax (Linaria cavanillesii)

Knowledge about the reproductive system of species inhabiting rocky habitats is scarce. The reproductive biology (floral biology, experimental pollination, insect visits, inbreeding depression, and seed predation) of the rupicolous Linaria cavanillesii was analyzed under field and experimental conditions. Self-compatibility was revealed by the high fruit set, seed set, and seed mass in the pollination experiment. Furthermore, results disclose that this species does not need insect visitors for seed production since fruit set after autonomous self-pollination was similar to that by hand cross-pollination. Self-offsprings were not affected by a strong degree of inbreeding depression in early life-cycle stages. Seed predation by Cucurlionidae is the main limiting factor of fruit and seed production in this species. Contrary to other perennial species of Linaria previously studied, L. cavanillesii represents the first clear report of self-fertility.     

Effect of gibberellic acid- and water-based pre-soaking treatments under different temperatures and photoperiods on the seed germination of Allium stracheyi Baker: An endangered alpine species of Central Himalaya,India

Allium stracheyi Baker (Alliaceae, 2600–3000 m asl), an endangered species of Central Himalaya, India, has low seed germination in its natural habitat. This study is an attempt to improve seed germination by determining the seed viability with a low mean germination time (MGT) and germination index (GI) under optimum temperature, light, and pre-soaking treatments. The seeds were pre-soaked in hot water (80°C), cold water (10°C), and gibberellic acid (GA₃ at 50 and 100 mg/l) for 24 h and subjected to light (12 h light and 12 h dark) and continuous dark (24 h) conditions with different temperature regimes (10, 15, 20, 25, and 30°C). The viability varied between 66.0% and 69.67% and declined rapidly after 12 months of storage. Our studies suggest that the 100 mg/l GA₃ treatment was beneficial for seed germination and seedling growth. Pre-soaking in a 100 mg/l GA₃ solution and incubation at 20°C under light conditions enhanced the germination significantly (p < 0.05) and resulted in the highest (97.3%) germination with the lowest MGT = 5.7 days, with GI = 8.11. The recommendations of this study support the conservation of alpine A. stracheyi via simple and cost-effective techniques for optimal seed germination.     

Multiple causes of seedling rarity in scrub plum, Prunus geniculata (Rosaceae), an endangered shrub of the Florida scrub

Conservation of an imperiled plant often requires an understanding of its reproductive ecology. Scrub plum (Prunus geniculata) is an endangered Florida shrub endemic to pyrogenic xeric uplands. Although plants are long-lived and may flower profusely, particularly after burning, fruit yield is sparse and seedlings are rare. We investigated potential causes of seedling rarity in scrub plum by studying its floral sex and breeding systems, fruit development, and germination ecology. We also developed a flow chart based on two time-since-fire scenarios to illustrate the cumulative impact of these factors on seedling recruitment. Breeding system experiments suggested partial gametophytic self-incompatibility in this functionally andromonoecious species, with inbreeding depression in self-compatible individuals. Predispersal seed predation, the major constraint on seedling recruitment, was lower in the first postburn season. In two field experiments, seed germination was <11% irrespective of treatment. These results indicate that seedling recruitment in scrub plum may be limited by multiple factors, including sexual reproductive failure, high predispersal seed predation, and low seed germination. The chance of a bisexual flower producing a seedling ranges from 5 in 1000 to 5 in 100000, depending on the time since fire. Restoration of scrub plum's historical fire regime may mitigate some of these factors.     

Implications of the reproductive biology of the narrow endemic <Emphasis Type="Italic">Naufraga balearica</Emphasis> (Apiaceae) for its conservation status

Naufraga balearica is an endemic umbelliferous plant of Mallorca (Balearic Islands, Spain), considered Critically Endangered according IUCN categories. We present a study of its reproductive biology to look for the causes of its threatened status. It was carried out in natural populations at Cap de Catalunya and Coves Blanques and in ex-situ cultivated plants on the campus of the Universitat de les Illes Balears. We examined (1) the flowering and fruiting phenology, reproductive potential and reproductive success; (2) the pollen/ovule (P/O) ratio and mating system; (3) the identities and numbers of pollinators; (4) the germination potential and the effect of temperature on the germination process; and (5) the seedling performance of this species. The results indicated that under natural conditions, an average individual produced approximately 100 seeds and cross-pollination was the most effective mating system. Pollination was myrmecophilous, and the germination potential was high (76–97%), but seedling survival after the first summer was low (7–54%). We conclude that the major threats to this species are fundamentally extrinsic in character and are related to changing climatic conditions, soil erosion and alteration, and umbel predation. Given that cross-pollination yields greater reproductive success and that pollination is myrmecophilous, lower population densities would lead to lower reproductive success for the species. Therefore, conservation efforts should focus primarily on habitat conservation and on ex-situ conservation through the establishment of an ex-situ population and the maintenance of a seed bank.     

Floral display and habitat fragmentation: Effects on the reproductive success of the threatened mass‐flowering Conospermum undulatum (Proteaceae)

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Unexpectedly high outcrossing rate in both dense and sparse patches in self-compatible Pedicularis rex (Orobanchaceae)

Pedicularis rex, a self-compatible alpine herb, depends exclusively on pollinators for reproduction. Previous study has revealed that 74.7 % of genetic variability occurs among populations in Yunnan, indicating a mixed mating system for the species. The large floral display also indicates high potential geitonogamy. Therefore, plant density and display size may be important factors influencing the realized mating system of P. rex. In the study reported here we explored the effects of plant density and floral display size on pollinator movements and thus outcrossing rate in two patches of in P. rex at low and high plant densities. We examined foraging behavior of bumblebees, measured reproductive outputs under hand and natural pollination, and estimated the outcrossing rates using RAPD markers. Floral display size was significantly larger and bumblebees visited significantly more flowers in sequence on individual plants in the sparse patch than in the dense patch. We found a significant positive correlation between floral display size and successive visits in the sparse patch, but not in the dense patch. Regression analysis revealed a significant inverse correlation between fruit production and flower production per plant with a theoretical maximum of 63 fruits per plant, but no relationship between seed production and flower production per plant. Contrary to previous studies on genetic structure, however, we found an unexpectedly high outcrossing rate in both the sparse and the dense patch (t _m = 1.151 and 0.924, respectively). Resource limitation may intensify seed competition and result in selected seed abortion, which may largely explain the unexpectedly high outcrossing rate.     

Prediction of cotton seedling germination against pre-emergence damping off on the basis of environmental factors and seed applied fungicides

Cotton is an important fibre crop chiefly attacked by a number of diseases. Pre-emergence damping off seedling is an important disease of cotton. Efficacy of three fungicides (Dynesty, Topsin-M and Antracol) was assessed on early mid and late sowings. Topsin-M and Antracol were evaluated at 1X and 2X concentrations, Dynasty was only used at standard dose. Significant increase in seed germination was seen, maximum germination was observed in late sowing. Antracol (2X) significantly improved seed germination rate in all three sowings as compared to other treatments. Impact of environmental parameters on germination rate was assessed relating to seed dressing fungicides at 1X and 2X concentrations. Significant correlation was observed between soil temperature and seedling germination rate by all treatments, 30–31 °C favoured germination rate. Maximum seedling germination rate was observed at 24–26 and 22–23 °C and 45–50% by average air temperature, minimum air temperature and relative humidity respectively. Regression model was contrived to predict seedling germinations impacted by seed dressing fungicides relating to environmental factors. Closed resemblance between observed and predicted values states that model can be used to predict seed germination by treating with fungicides regarding to environmental factors.     

Reproductive characteristics that favor invasiveness in Leonotis nepetifolia (L.) R. Br.

Reproductive systems are life attributes important in defining the demography and genetic constitution of invasive alien species populations. We describe the phenology, floral behavior and floral visitors in Mexican populations of Leonotis nepetifolia considered invasive in America, Asia and Oceania. The mating system was determined through pollination experiments and, with a morphological analysis of flowers (outcrossing index, OCI) and pollen/ovule ratio, the breeding system was evaluated. Germination of 1 and 2-year-old seeds was tested to assess the potential characteristics of germination. Leonotis nepetifolia was reproductive for 7 months (June to December) and tended towards a specific season during autumn. Anthesis lasted 36 hr with protogyny and no hercogamy, with floral visitors of Apodiformes, Hymenoptera, Lepidoptera and Thysanoptera. Pollination experiments indicated a mixed mating system, whereas the OCI and the pollen/ovule ratio pointed towards a facultative xenogamous breeding system. Seed production was high (1,445 ± 132 seeds/plant); the seeds had potential longevity and were neutral photoblastic. One-year-old seeds germinated slightly later ( = 2.6 ± 0.11 days) than 2-year-old seeds ( = 1.9 ± 0.02 days), both synchronously (IS_1yr = 0.88 ± 0.03 and IS_2yr = 0.82 ± 0.02). Germination percentage for 1-year-old seeds was lower (55.33 ± 4.40%) than that of 2-year-old seeds (94.18 ± 0.59%), suggesting a potential longevity of the seeds in an optimal environment. Reproductive characteristics, such as wide reproductive period, mixed breeding system, copious seed production, seeds with potential longevity, and quick and synchronic germination in different light conditions, favor the invasive capacity of Leonotis nepetifolia.     

交配距离与父本数量对野慈姑有性繁殖的影响

植物的有性繁殖是生活史和进化的核心, 母本及环境对繁殖的影响固然重要, 父本的贡献亦不容忽视。父本来源与多样性对坐果结籽和后代质量的影响明显, 但由于不同物种或种群的繁殖特点和进化历程, 往往也会呈现其独特性。该研究旨在探究挺水植物野慈姑(Sagittaria trifolia)是否存在自交或远交衰退, 以及父本数量对其繁殖和后代表现的影响。通过人工控制授粉, 设置自交、近距离异交(<50 km)、远距离异交(＞200 km) 3种交配距离以及单、双两类父本数量, 共计5种授粉处理, 观测野慈姑坐果概率、单果种子数量、种子面积、萌发率(2018和2019年)、幼苗芽长(2018和2019年)共7项指标。结果显示: 不同距离的交配对野慈姑的坐果概率、单果种子数量、种子面积、萌发率、幼苗芽长没有显著影响, 野慈姑未出现明显的自交衰退或远交衰退现象。父本数量的增加对野慈姑的结实数量(坐果概率、单果种子数量、种子面积)无影响, 但结实质量方面, 双父本处理的种子萌发率略高于单父本处理。综上所述, 交配距离与父本数量对野慈姑的繁殖表现影响较小, 这可能与其频繁的自交历史、远距离的基因扩散以及高度相似的水生环境有关; 同时, 该研究基于20余个自然种群的基因型, 研究结果也表明了野慈姑物种水平有性繁殖的优异和稳定性。     

Reproductive versatility in legumes: the case of amphicarpy in Trifolium polymorphum

Amphicarpy is a fascinating reproductive strategy, defined as fruit produced both below the soil surface and as aerial fruit on the same plant. Trifolium polymorphum is a grassland species subject to herbivory that combines amphicarpy with vegetative reproduction through stolons. Underground flowers have been described as obligate autogamous and aerial ones as self‐compatible allogamous, with aerial floral traits favouring cross‐pollination. In the present work we performed different pollination treatments on aerial flowers to analyse rates of pollen tube development and offspring fitness, measured as fruit set, seed production and germination percentage. This last variable was compared to that of seeds produced underground. No significant differences were found between fruit set in self‐ and cross‐pollinations. Seed production was higher in self‐pollinations, which is consistent with the higher rate of pollen tube development observed in self‐crosses. Spontaneous self‐pollination is limited in aerial flowers; thus pollen transfer by means of a vector is required even within the same flower. Germination tests showed that aerial seeds produced after self‐ and cross‐pollination did not differ in fitness, but underground seeds had higher germination percentage than aerial ones. Thus, we conclude that T. polymorphum has a mixed mating system. In grasslands with heavy grazing pressure, clonal propagation and underground seed production ensure persistence in the field. An intermediate level of selfing in aerial flowers ensures offspring, but morphological (herkogamy) and functional (dicogamy) floral traits maintain a window to incorporate genetic variability, allowing the species to tolerate temporal and spatial pressures.     

Reproductive strategies of diploid and polyploid populations of arcticDraba (Brassicaceae)

132 cultivated populations (2x–16x) of 15 arctic-alpine species ofDraba were investigated to clarify a possible relationship between reproductive strategies and polyploid evolution in the genus. The populations were exclusively sexual and produced viable seed after spontaneous self-pollination, but showed large variation both in traits promoting cross-pollination and in autogamous fruit and seed set. Traits promoting cross-pollination, e.g., floral display, protogyny, and delayed selfing, were positively correlated, and these traits were negatively correlated with autogamous fruit and seed set. All diploid and many polyploid populations had high autogamous seed set and small, unscented, non-protogynous, and rapidly selfing flowers. In contrast, all populations with low autogamous seed set and large, scented, and strongly protogynous flowers with distinctly delayed selfing were polyploid. These results are consistent with those previously obtained from enzyme electrophoresis, suggesting that the genetically depauperate diploids are extreme inbreeders and that the highly fixed-heterozygous polyploids vary from extreme inbreeders to mixed maters. The reproductive data lend additional support to the hypothesis that allopolyploidy in arcticDraba serves as an escape from genetic depauperation caused by uniparental inbreeding at the diploid level.     

Reproductive biological characteristics potentially contributed to invasiveness in an alien invasive plant Bidens frondosa

Bidens frondosa L. (Asteraceae) is a widespread invasive weed in China. By experimental observation and bagging treatment, the reproductive biological characteristics of this species, such as phenology of flowering, floral syndrome, breeding system and seed germination characteristics, were studied to assess the association of these reproductive characteristics with invasiveness. Flowers of B. frondosa bloom from September to October every year in Ji'an city, Jiangxi province. The lifespan of a single capitulum is approximately 4 to 5 days, with 30 to 60 florets per capitulum. The capitulum diameter, anthocaulus length, floret length and width, stamen length and pistil length were 6.1, 30.9, 2.2, 0.6, 3.0 and 2.7 mm, respectively. The seed set percentage of 48.5% in the treatment of bagging flowers without emasculation suggests B. frondosa is self‐compatible; meanwhile, the percentage of 63.1% in the treatment of bagging with emasculation and manual xenogamy suggests it also is cross‐compatible. P/O ratio per capitulum of this species was 450.5, which suggests that the breeding system of B. frondosa is facultative xenogamous, and it needs pollinators to some degree. The main floral visitors were insects of Hymenoptera, Diptera and Lepidoptera. The 1000‐achenes weight was 1.97 g. The achenes emerged as a small germination peak from the fourth to fifth day, and had a high accumulated germination rate of up to 84.0% on day 20 after sowing. Our experimental findings suggested that the reproductive biological characteristics, such as the versatile mating system of self‐ and cross‐pollination, high seed production, a special method of achene dispersal and germination peak, accompanied by a high accumulated germination rate, might contribute to the invasive ability of B. frondosa.     

The avoidance of self-interference in the Tuscan endemic spring geophyte Crocus etruscus Parl. (Iridaceae)

Spatial separation between sexes within hermaphrodite flowers (herkogamy) is widespread among angiosperms and is traditionally viewed as an adaptation that reduces the likelihood of self-pollination. While different degrees of herkogamy have been reported for Crocus, the relationship between sexual organ positions and reproductive fitness has not been explored yet, and studies of the breeding system within natural populations remain scarce within the genus. We analyzed the effects of different breeding systems (spontaneous self-pollination, facilitated self-pollination and cross-pollination) and of herkogamy degree on female reproductive success of the Tuscan endemic Crocus etruscus. Results showed that C. etruscus is monomorphic for style length and self-incompatible (SCI = 0.29). We also detected a significant negative effect of herkogamy on seed set of open-pollinated flowers. Thus, we conclude that the species avoids self-fertilization by self-incompatibility, but interestingly shows also a certain degree of spatial separation of male and female reproductive structures within the same flower. This floral trait could be useful to avoid self-interference thus reducing ovule discounting and pollen loss on its own stigma.     

Long-term progression of inbreeding depression in a Mediterranean ornithophilous shrub

The ornithophilous species Anagyris foetida L. is a Mediterranean shrub with highly fragmented populations and a mixed mating system. In a previous study, we analyzed the first 3 years of the life cycle of two progenies (selfed and outcrossed) grown from seed obtained by hand pollination and planted in an experimental garden in 2005. In that study, we found that inbreeding depression (ID) was manifested both reproductively and vegetatively throughout the life cycle, with male reproductive function being the most affected trait. In the present study, our main aim was to check the progression of the two progenies 12 years after transplantation. For this we analyzed their survival and their vegetative and reproductive traits. According to our results, levels of ID were similar to those obtained in the previous study, with some factors decreased and thus varying with the age of the studied individuals. Vegetative parameters were found to have a greater influence than reproductive ones (δ = 0.56 vs. δ = 0.36) on overall ID. As indicated by the global ID (δ = 0.72) the populations have a mating system that is intermediate between outcrossing and a mixed system. Furthermore, the lower male reproductive capacity of the selfed individuals has been maintained over time. Our study also demonstrates the importance of studying the ID value throughout the life cycle of plants.     

菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系

三叶鬼针草(Bidens pilosa)是一种危害严重的菊科入侵植物。通过实验观察和人工控制套袋等方法, 对其花序开花动态、花粉胚珠比 (P/O)、自交亲和性、花粉活力、访花昆虫和种子 (瘦果)的萌发率等与繁殖相关的特征进行了研究, 探讨了这些繁殖特征与入侵性的关系。三叶鬼针草在10-11月开花, 单个花序的花期约为5-6天。每小花的花柱基部有圆筒状的蜜腺环绕。单个花序内可自交亲和, 自交结实率和花粉活力均较高, 其P/O值为1 7 54.1 2±29 .8 7。主要访花者为灰蝶科(Lycaenidae)、粉蝶科(Pieridae)和茧蜂科(Braconidae)昆虫。三叶鬼针草所具有的灵活交配机制是其入侵成功的重要因素。此外, 三叶鬼针草结实量大、种子产生迅速且适于传播, 以及种子萌发范围广和短期快速萌发等特性也增强了其入侵性。     

Laboratory evaluation of grain protectant efficacy of different foliar extracts against maize weevil – Sitophilus zeamais (Motschulsky)

The present study was aimed to evaluate six indigenous plant extracts for their ability to protect maize from infestation by maize weevil Sitophilus zeamais (M). The plant extracts were tested for its toxicity by fumigation and grain surface coating methods. Further, the effects of botanical treatments on F1 progeny, seed emergence and antifungal activities were studied. The results indicated that the extracts of Eicchornia crassipes controlled the test insect in grain surface coating method (100%, P = 0.05). Further, the same plant extract had significantly reduced the development of F1 progeny (97 ± 0.7%). In germination experiments, T + UEx samples showed normal seed germination of maize. However, in T + Ex samples, the seed germination depended on the grain protectant efficacy of the tested botanical extracts. Always higher percantage seed germination was recorded in those treatments where there was less infestation. Extracts of Carica papaya had significantly reduced the incidence of seed-borne fungus Aspergillus flavus (4 ± 2.4%) and did not affect the seed germination when compared to other treatments and untreated control.     

Abnormalities in sexual development and pollinator limitation in Michelia coriacea (Magnoliaceae), a critically endangered endemic to Southeast Yunnan,China

Michelia coriacea (Magnoliacae) is a critically endangered tree, endemic to Southeast Yunnan province, China. Most of the individuals in the extant populations normally bear flowers, but fruit set and fertile seed production rates were only 6.7% and 0.2%, respectively. To identify possible causes of reproductive barriers, the following studies were carried out: examination of stamen and pistil development; experiments using controlled pollination; observations of behavior and frequency of floral visitors in both cultivated and natural populations. The results revealed that about 60% of pollen was abnormal and approx. 70% of ovules had delayed development. Hand pollination could effectively enhance the fruit set (F_4,25=35.139, P<0.0001) and seed set per fruit (F_4,25=85.022, P<0.0001). Both cultivated and wild M. coriacea had an extremely low frequency of floral visitors. Some beetles, a few species of Andrenidae and some Bombus sp. are likely to be the effective pollinators. The fruit set and seed set per fruit from controlled self-pollination and cross-pollination were significantly different (P<0.05), and thus it is inferred, that inbreeding depression may be a contributing factor in the very low seed production. It appears that low seed set in M. coriacea is due to a combination of factors: abnormalities in pollen and ovules, low number of effective pollinators, and inbreeding depression.     

Pollination biology of Aristolochia bianorii Sennen & Pau: promoting cross-pollination but assuring reproductive success in island ecosystems

• Deceptive pollination has been reported in the genus Aristolochia, but the floral biology and pollination strategy of A. bianorii, an endemic of the Balearic Islands, have not yet been studied. Here, we investigated floral anthesis, mating system, pollinators and volatile organic compounds (VOCs) emitted by its flowers.
• Flower buds were marked and monitored daily to define floral stages and their duration. Experimental bagging and hand-pollination were performed to test for autonomous self-pollination, induced self-pollination and cross-pollination. Flowers were collected to analyse the presence of entrapped pollinators. VOCs emitted by flowers were evaluated by means of solid phase microextraction followed by immediate GC–MS.
• Anthesis lasted between 63 and 96 h, and the species exhibited autonomous self-pollination with moderate inbreeding depression. Pollinators were mainly females of Oscinomorpha longirostris (Diptera; Chloropidae). The number of pollinators inside flowers was affected by floral stage and time of flowering. The most common VOCs were alkanes, oximes, esters, alkenes, cyclic unsaturated hydrocarbons, isocyanates, amides and carboxylic acids.
• Aristolochia bianorii can set seed by autonomous self-pollination, in contrast to other Aristolochia species, in which both protogyny and herkogamy prevent autonomous self-pollination. However, the species may encourage cross-pollination by attracting female chloropid flies though emission of floral scents that may mimic an oviposition site and, possibly, freshly killed true bugs (i.e. Heteroptera). In conclusion, A. bianorii promotes cross-pollination, but delayed autonomous self-pollination assures reproductive success in the putative absence of pollinators.

     

Floral biology and potential hybridization of three sympatric epiphytic bromeliads in Veracruz,Mexico

In the Bromeliaceae, hybrids have been engineered for horticultural purposes; in addition, hybrids have been observed in natural habitats. We investigated the hybridization potential of three sympatric species of Tillandsia inhabiting secondary forests and shaded coffee plantations in the central mountains of Veracruz, Mexico. The studied species (T. juncea, T. polystachia and T. variabilis) show overlap in flowering phenology and are believed to share pollinators. We compared the floral morphology of the species, performed controlled intra- and interspecific pollinations, and recorded pollen tube growth, fruit and seed set, as well as seed germination. Flower size differed among the species, but stamen and stigma heights were similar between pairs of species, which would facilitate interspecific pollen exchange. Pollen tubes reached the ovary in interspecific crosses. The three species showed interspecific cross-compatibility, as all crosses resulted in development of mature fruit and viable seeds. Tillandsia juncea achieved the highest fruit set, whereas T. variabilis showed the highest seed set. Seed set as a result of agamospermy, autogamy, geitonogamy and interspecific cross-pollination did not differ among the species. The findings suggest that pre- and post-pollination barriers are weak among the three species of Tillandsia, which thus could potentially hybridize in natural sympatric populations.