Vivipary in Ophiorrhiza mungos L. – a rare phenomenon in angiosperms

Vivipary, the precocious germination of seeds within the parent plant, is a specialised feature of evolutionary and biological importance that ensures survival of a plant. Reports on vivipary in angiosperms are rare, accounting for <0.1% of flowering plants. Here, we report a remarkable case of occurrence of vivipary in Ophiorrhiza mungos. A study was conducted to collect information on the morphology of the capsules that support vivipary, environmental factors that induce vivipary, survival mode and the survival of viviparous seedlings. The hydroscopic movement of the cup‐shaped capsules of O. mungos was found to help in viviparous germination during the rainy season. Of the total seeds in a capsule, 70% showed viviparous germination. The seedlings remaining inside the capsule attain a height of 0.98 ± 0.4 cm and reach the ground when the capsule falls. On the ground, seedlings obtain easy anchorage to the substratum since they have already germinated. Vivipary appears to be an adaptation of O. mungos to the rainy season for ensuring viable offspring. This suggests that vivipary in this species might be artificially induced by continuous spraying with water to rescue seeds in all seasons for use in large‐scale propagation to meet increasing market demand and conservation of this valuable anticancer medicinal herb.     

Vivipary and offspring survival in the epiphytic cactus Epiphyllum phyllanthus (Cactaceae)

Vivipary, the germination of seeds before they are shed from the parent plant, is a rare event in angiosperms involving complex ecophysiological processes. Pseudovivipary and cryptovivipary occur in approximately 30 (2%) species of the cactus family. A remarkable case of vivipary in Epiphyllum phyllanthus is described here. Information is provided regarding the biology of viviparous fruits, morphology, mortality, survival rates of viviparous offspring, and some eco-evolutionary implications of this reproductive strategy in the Cactaceae. This epiphytic cactus has no host-specific relationship. A low proportion (33.3%) of individuals produced viviparous fruits. Seed number/fruit varied from 197 to 230 with percentage of viviparous germination from 97.5% to 99%. The viviparous seedlings exhibited normal development and were no different from non-viviparous offspring. Transplanting experiments showed that the first week is critical for seedling establishment, and high mortality occurred in the three treatments used: 69% on the phorophyte surface, 58.6% on the ground, and 44.8% under controlled conditions. The number of survivors gradually stabilized, and the contribution to establishment was comparable in each of the treatments after the acclimation phase. It is suggested that vivipary is associated with thermoregulation, parental care, conspecific nursing, and rapid seedling establishment. Germination is not a limiting factor in the perpetuation of this viviparous species, but seedling establishment is. In viviparous individuals of E. phyllanthus, seedling mortality during establishment rather than failure to germinate within the fruit is a limiting factor affecting local population density. Overall, viviparity is an intrinsic, genetic event involving high metabolic costs favouring germination and dispersal of the fittest offspring regardless of substrate and environmental conditions.     

Vivipary in the Cactaceae: Its taxonomic occurrence and biological significance

Vivipary, a rare reproductive strategy in flowering plants, has been previously reported in only four species of the Cactaceae. In this paper, eight additional examples (Cleistocactus smaragdiflorus, Disocactus martianus, the hybrid Epiphyllum X Fern la Borde, Harrisia martinii, Lepismium ianthothele, L. monacanthum, Rhipsalis micrantha forma micrantha, and R. baccifera subsp. horrida) of vivipary in cultivated cacti from tribes Hylocereeae, Rhipsalideae, and Trichocereeae are documented. The plants had fully-developed and mature fruits containing seeds at different stages of germination, including tiny embryos emerging from the seed coat to young, healthy seedlings with well-developed organs, such as cotyledons and radicles. Several features, though not unique, distinguish cactus fruits with viviparous seedlings. Foremost, the fruits are fleshy with abundant mucilage and a relatively thick, often transparent pericarp. The seeds are generally numerous and embedded in a transparent, white or red pulp. The occurrence of vivipary in taxa from four phylogenetically distinct tribes of the subfamily Cactoideae suggests an independent origin and indicates that this reproductive strategy may be more widespread in the family than originally thought. Vivipary in the Cactaceae is regarded as a form of cryptovivipary, a condition in which the zygote develops inside the fruit without penetrating the ovary wall for dispersal purposes, which in turn is considered a subcategory of true vivipary. Further, vivipary is interpreted as an adaptive reproductive strategy that enables seedlings to establish more rapidly.     

植物营养体胎生研究进展

营养体胎生是植物体上的营养器官如胞芽、珠芽、不定芽、叶片等成熟后不脱落,直接吸取母体营养继续生长发育,在自然状态下脱离形成新的植株。营养体胎生植物的种类较多,大多是在长期适应干旱、冷凉、高温等恶劣环境条件下进化出的结果,是繁育途径的一个重要补充。与传统的无性繁殖方式相比,繁殖系数较高,可实现高效再生,且病毒感染率较低,在实际生产中具有重要的应用价值。从胎生器官的形态发育、生理生化、分子水平和再生繁殖特性4个方面进行了综述,目前研究还处于起步阶段,对引起胎生现象的本质还没有得到系统解答,指出存在的不足,并对未来的研究方向进行了展望,关于营养体胎生形成机理的调控和重要功能基因的挖掘,以及胎生苗如何适应新的环境变化也是未来的研究方向。因此,研究营养体胎生适应性现象的本质及其进化将为植物种群的繁衍、规模化生产和应用、生物多样性保护等提供理论依据。     

Bidens pilosa L.: Exclusive report of vivipary in a non-mangrove taxa from the eastern Himalayas

Vivipary is an extremely evolved type of epigeal germination in seed plants and very common in tropical estuarine plant communities (mangroves), ensuring a greater number of viable settings of offspring. In addition to this, there are some other non-mangrove angiosperm taxa that have a successful existence, and because of their invasive nature, vivipary has been reported as an attribute of their uncongenial abiotic stress. The present observation confirms the incidence of true vivipary in Bidens pilosa L. (a member of the family Asteraceae) from the eastern Himalayan region.     

红树植物胎生现象研究进展

植物胎生是指有性繁殖产生的后代在母体上直接萌发的现象, 在红树植物中最为常见。红树植物生长在热带亚热带海岸潮间带, 耐受高盐、高温、淹水缺氧和海浪冲击等复杂环境。胎生被认为是红树植物对这种特殊生境的重要适应方式。该文从形态发育、生理生化、分子水平、生态适应4个层次讨论红树植物胎生现象对复杂生境的适应性, 并指出现有研究存在的不足, 对将来的研究方向进行了展望。与非胎生胚胎发育相比, 红树植物胎生是一个遗传的程序, 在进化过程中形成了一些特殊的结构。植物激素对胎生发育起关键的调控作用, 繁殖体发育过程中, 其盐离子的种类与浓度的动态变化则是对海岸潮间带生境的重要适应特征。这种胎生繁殖体依靠在母体上完善的一系列功能性特征能更有效地适应落地后的滩涂环境。然而, 红树植物胎生发育过程的分子机理及调控机制还有待研究。理解胎生这一特殊适应性现象的本质及其进化过程将为红树林保护繁育、适应气候变化提供理论依据。     

Progress in the studies of vivipary in mangrove plants

Vivipary in plants refers to a phenomenon that sexually reproduced offsprings germinate while still attached to the maternal bodies. This is mostly manifested in mangrove plants, which occur in tropical and subtropical intertidal zones and encounter harsh environmental conditions such as high salinity, high temperatures, waterlogging, hypoxia and tidal waves. Vivipary has long been recognized as one of the most important adaptive features under such a complex environment. Here we discuss four aspects of vivipary: morphological anatomy, physiology and biochemistry, molecular biology and ecological adaptation. We also discuss shortcomings in current studies and prospect of future directions. Differing from regular seed development, viviparous seeds in mangroves are evolved with many special structures, indicating a genetically based process. Hormones play an important role in regulating the process, whilst the dynamics of salt ion concentration during embryo and propagule development seems to be an adaptive feature. The ecological significance of vivipary is fully exhibited in the propagules that can effectively establish themselves on muddy tidal zones. Such a success heavily relies on sound functional features developed on the mother plants. However, the molecular mechanism and the regulation of viviparous seed development in mangroves remain elusive. Systematic studies of vivipary in mangroves not only help to understand the nature and evolutionary process of this distinct adaptive phenomenon, but also provide the foundation for mangrove forest restoration and protection in many parts of the world.     

Interspecific Variations in Responses of Mangrove Seedlings to Two Contrasting Salinities

The growth performance of seedlings of seven species of true mangroves (Avicennia marina, A. officinalis, Bruguiera gymnorrhiza, B. sexangula, Rhizophora apiculata, R. mucronata, and Sonneratia caseolaris) in response to two contrasting salinity regimes, low (i.e., 3–5) and medium (i.e., 25–27), was studied. Species represented all categories relevant to vivipary (i.e., true viviparous species, cryptoviviparous species and non‐viviparous species), and included closely related pairs as well as species commonly used in replanting in Sri Lanka. Species could be ranked in descending order of salinity tolerance as A. marina > R. mucronata > R. apiculata >B. gymnorrhiza > A. officinalis > B. sexangula > S. caseolaris, hence taxonomically similar species and those sharing vivipary characteristics may be distant in salinity tolerance. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Dormancy release, ABA and pre-harvest sprouting

Seed dormancy is an adaptive trait that enables the seeds of many species to remain quiescent until conditions become favorable for germination. Dormancy is normally initiated during seed maturation and maintained to seed maturity. In mature seeds, the loss of dormancy may be gradual (after-ripening) or can be terminated by chilling and other environmental triggers. Dormancy is an important trait for many important crop species: it inhibits pre-harvest spouting or vivipary, a widespread problem in many regions of the world. Too much dormancy, however, can lead to non-uniform germination in the field. Recent progress has been made in understanding the role of abscisic acid metabolism and dormancy release in both model plants and crop species. Advances in our understanding of the molecular mechanisms that are involved in dormancy, along with approaches using quantitative genetics, will provide new strategies through which the desired level of dormancy can be introduced into crop species.     

Can germination explain the distribution of tree species in a savanna wetland?

The relationship between seed germination and ecological niche is determined by matching germination characteristics with environmental features. In this study, we selected tree species occurring in the largest savanna wetland in South America – the Pantanal. Very few species are endemic or exclusively found in savanna wetlands, and the majority of tree species occurring in the Pantanal are also found in the neighbouring Brazilian Cerrado, a drier vegetation type that does not flood. We investigated the relationship between germination characteristics and occurrence of savanna trees in wetlands testing the hypothesis that such seeds are tolerant to flooding. We also addressed the question of whether seed tolerance to flood, assessed by survival analysis, explains tree distribution along a gradient of flooding intensity. In this flooding gradient, widely distributed species are those that occur in areas subjected to low as well as to high flooding intensity whereas restricted distributed species are those that occur only in areas subjected to a low level of flood. Seeds from tree species occurring in areas subjected to different flooding intensities were collected. Seed tolerance and germination during and after both one and two months of simulated flood were evaluated. Our results show that seeds of most of the studied savanna species tolerated submergence, which helps to explain their occurrence and wide distribution in wetlands. Nevertheless, germination behaviour checked by survival functions (i.e. how germination is distributed over time) partially explained tree species distribution along a flooding gradient. We conclude that seed tolerance to flooding is one of the components of the regeneration niche that determines tree occurrence and distribution at the regional scale, from savanna to wetland, but not at a local scale along a flooding gradient.     

Flooding tolerance of Tabebuia cassinoides: Metabolic, morphological and growth responses

Tabebuia cassinoides (Lam.) DC (Bignoniaceae) is a tree species that occurs in swampy areas of the coastal “restinga” in SE Brazil (a coastal sandy plains scrub and forest formation). To elucidate possible adaptive strategies that enable this species to occupy areas subjected to seasonal or perennial waterlogging, metabolic, morphological and growth responses of plants under flooding conditions were studied. The root system of T. cassinoides plants presented elevated amounts of ethanol (10.6 μmol g⁻¹ fresh wt) only in the first 5 d of soil water saturation. The two-fold increase in ethanol production under flooding was corroborated by an increase in ADH activity in the same period. Lactic acid concentrations did not change significantly during four months of flooding treatment. The decrease of alcoholic fermentation under hypoxia was associated with the appearing of new roots. The induction of aerenchyma formation in roots developed under flooding conditions, allowed oxygen transport from the shoot to these organs, thus maintaining an aerobic respiration. We conclude that this characteristic and the capacity to oxidize the rhizosphere are probably responsible for the survival and growth of plants while flooded and for their success in an environment, which restricts the presence of the majority of competing tree species.     

Reductions in abscisic acid are linked with viviparous reproduction in mangroves

We investigate physiological mechanisms behind the convergent evolutionary loss of seed dormancy in plant lineages, focusing on mangroves as a model system. More than 60 angiosperm families, including several mangrove taxa, contain species with seeds that are intolerant of drying and do not undergo dormancy. These desiccation-intolerant species occur with disproportionate frequency in wet or coastal tropical habitats. In plants, the hormone abscisic acid (ABA) coordinates both the development of desiccation tolerance during the onset of seed dormancy and whole-organism responses to flooding. Thus, changes in ABA levels and/or modes of action in different plant compartments are implicated in the repeated evolutionary loss of seed dormancy among species of wet habitats. We compare ontogenetic dynamics of ABA levels in embryonic, maternal, and mature vegetative tissue of four phylogenetically independent pairs of related viviparous mangroves and nonviviparous nonmangroves. We demonstrate that ABA levels are consistently lower in embryos of viviparous mangrove taxa than embryos of nonmangrove, nonviviparous sister taxa. In contrast, elevated tissue concentrations of ABA characterize leaves of all mangrove species tested, while ABA levels in maternal tissues vary among mangrove species. These commonalities suggest a functionally important trade-off between the maintenance of embryonic development and the adjustment of the parent tree to salinity stress. This study yields comparative data on seed physiology in naturally occurring desiccation-intolerant species, for which these data are currently scarce, and demonstrates a potentially significant role of phytohormones in the evolution of plant life histories.     

Vivipary as a tool to analyze late embryogenic events in maize

In maize vivipary, the precocious germination of the seed while still attached to the ear, is the diagnostic phenotype of mutants, which are impaired in the biosynthesis or response to abscisic acid (ABA). Of the 15 genes so far described, 12 control specific steps in ABA biosynthesis, two mediate hormone response and one still has an undefined role. We have analyzed a collection of 25 independent vp isolates with the aim of determining the degree of mutational saturation that has so far been reached. Of the 25 viviparous mutants complementation tested, 22 correspond to known loci: six are allelic to vp1, another six to vp5, one to vp7, two to vp9, six to vp10 and one to w3. The remaining three represent genes not previously identified. All mutants so far tested except rea show a decrease in ABA content. As to the only two mutants (vp1 and rea) whose endogenous ABA content is not impaired, the reduction in sensitivity of the double mutant compared to the single ones suggests that the two genes control separate pathways in the ABA signal transduction. Some of the mutants in this collection have a characteristic incomplete germination that allows the embryo of the mature dry seed to resume germination. By exploiting this feature it is possible to infer, through a germination test, whether the mutant has been impaired in the acquisition of desiccation tolerance. This information provides the starting point for the dissection of the genetic basis of desiccation tolerance.     

Contributo Allo Studio Della Temperatura Delle Infiorescenze Delle Palme E Aracee

Abstract

Prioritization of endemic, threatened species and the habitats where they live is a crucial point of conservation actions, particularly in areas with rich endemic floras. In this study, we have developed a new procedure to define the conservation priorities among endemic plants and habitats by evaluating eight criteria. Five criteria deal with the geographic and ecological range of the evaluated species, whereas the other three refer to threats. After the evaluation of each criterion, we combined the partial scores to obtain a priority index (PI). Finally, we characterized the EU habitat categories of conservation concern on the basis of the average PI value of the plants living in each habitat. We tested the method on a list of 260 endemic plants from a biodiversity hotspot (Sardinia) that had an average PI of 3.66 ± 0.16. Even if the habitat categories that are most rich in endemic plants were rocky habitats, and coastal/halophytic habitats, the most endangered habitat was coastal sand dunes (PI = 6.75 ± 1.15). The method herein presented is complementary with the application of IUCN criteria. This integrated approach is a concrete solution that adapts IUCN criteria and categories to local contexts.     

Shingle habitats in the British Isles

RANDALL, R. E., 1989. Shingle habitats in the British Isles . Shingle is the term applied to sediments larger in diameter than sand but smaller than boulders. Around Great Britain there are almost 900 km of pure shingle and vast stretches of rock/shingle, sand/shingle and mud/shingle mixtures. Outside Japan, New Zealand and north-west Europe, shingle is an uncommon coastal sediment. Shingle may form fringing beaches, spits, bars, cuspate forelands and offshore barrier islands, depending upon available supply of sedimentary material and coastal topography.
Species composition on shingle features is partly determined by climate which affects the geographic range of certain species. The other major factors are beach stability and beach composition. Climate results in distinctive patterns in the flora but within the each climate zone beach movement will affect the quantity of annuals, short-lived or long-lived perennials present. The presence or absence and nature of the fine material within the shingle will cause different combinations of species ecologically related to those of sand dunes, salt-marsh or cliff. Coastal shingle vegetation has a distinctive flora which contains several rare or declining species as well as some common coastal and ruderal plants.
At the largest shingle sites a successional sequence can be recorded which initiates with halophytic forbs near the foreshore and moves through neutral to acid grassland species with shrubs and prostrate scrub in the more inland areas.     

The sessile oak: Anomalies of the binomial

Summary

Plantlets of viviparous grasses produce mature vegetative plants of greater stature than those from seeds of closely related seed-bearing grasses. Results from mixed populations of Festuca vivipara and F. ovina indicate size differences are maintained at flowering. Viable plantlets do not dehisce readily from mature panicles and when detached artificially require a large amount of free water to germinate. Results are discussed with reference to the occurrence of vivipary in Scottish grasses and its success in upland zones.     

Phylogeography of halophytes from European coastal and inland habitats

The repeated advance and retreat of glaciers during the Pleistocene ice ages have played a major role in shaping the present patterns of genetic variation within and among plant and animal populations of the temperate zone. In Europe, the geographic ranges of many species were confined to a few, mostly southern refugia during periods of full glaciation. Distribution ranges then reexpanded, and uninhabited northern areas were recolonized during the interglacials. These contraction–expansion cycles were repeated at least four times. Paleontological and molecular phylogeographic studies during the last decade have greatly increased our knowledge of refugial areas and postglacial recolonization patterns of European trees, shrubs and Alpine plants since the last glacial maximum about 20,000 years ago. Much less is known yet about non-Alpine herbaceous plants. In the present review, we summarize recent phylogeographic work on halophytic (salt-adapted) plants from coastal and inland habitats in Europe. Major refugial areas for these plants have been identified along the Mediterranean coasts, but some species could also have survived in saline inland localities. In general, recolonization of N and NW Europe occurred in a stepwise fashion along the Atlantic coastline. For a number of species, molecular studies revealed concordant genetic discontinuities on the background of an essentially continuous geographic distribution. Such congruency could be explained by the preferential seed dispersal through sea currents. However, phylogeographic patterns of halophytes also proved to be influenced by other factors like sea-level fluctuations during the Pleistocene, secondary contact between divergent lineages, long-distance dispersal, clonal growth, and special habitat and temperature requirements.     

Impacts of Carpobrotus edulis (L.) N.E.Br. on the Germination,Establishment and Survival of Native Plants: A Clue for Assessing Its Competitive Strength

Does Carpobrotus edulis have an impact on native plants? How do C. edulis’ soil residual effects affect the maintenance of native populations? What is the extent of interspecific competition in its invasion process? In order to answer those questions, we established pure and mixed cultures of native species and C. edulis on soil collected from invaded and native areas of Mediterranean coastal dunes in the Iberian Peninsula. We examined the impact of the invader on the germination, growth and survival of seeds and adult plants of two native plant species (Malcolmia littorea (L.) R.Br, and Scabiosa atropurpurea L.) growing with ramets or seeds of C. edulis. Residual effects of C. edulis on soils affected the germination process and early growth of native plants in different ways, depending on plant species and density. Interspecific competition significantly reduced the germination and early growth of native plants but this result was soil, density, timing and plant species dependent. Also, at any density of adult individuals of C. edulis, established native adult plants were not competitive. Moreover, ramets of C. edulis had a lethal effect on native plants, which died in a short period of time. Even the presence of C. edulis seedlings prevents the recruitment of native species. In conclusion, C. edulis have strong negative impacts on the germination, growth and survival of the native species M. littorea and S. atropurpurea. These impacts were highly depended on the development stages of native and invasive plants. Our findings are crucial for new strategies of biodiversity conservation in coastal habitats.     

Premature Drying, Fluridone-Treatment, and Embryo Isolation During Development of Maize Kernels (Zea mays L.) Induce Germination, but the Protein Synthetic Responses are Different. Potential Regulation of Germination and Protein Synthesis by Abscisic Acid

Freshly harvested, developing kernels of maize (Zea mays L.)do not germinate up to 77 d after pollination, but can be inducedto do so by fluridone, premature desiccation, and isolationof the developing embryo. The pattern of protein synthesis indeveloping maize embryos is distinct from that during germinationand subsequent seedling growth. Premature desiccation at 35DAP elicits a pattern of protein synthesis upon rehydrationwhich is similar to that in germinated embryos from mature drykernels. Fluridone-induced viviparous germination is accompaniedby changes in the synthesis of some proteins to a post-germinativepattern, but some developmental proteins continue to be synthesized.Embryos isolated from developing kernels at 35 DAP germinatewhen incubated on water; they also produce some developmentalproteins during germination. Kernels from developing cobs at35 DAP which are detached from the mother plant and maintainedin an atmosphere of high relative humidity (moist controls)do not germinate, but neither do they continue a clearly definedpattern of either developmental or germinative protein synthesis.Drying is thus critical to effect a clear transition of proteinsynthesis from a developmental to a germinative mode in maizeembryos. Abscisic acid within the developing embryos is reduced by fluridone,but to a lesser extent by premature drying or maturation drying.Changes in sensitivity to abscisic acid by the developing embryomay be as, or more, important in permitting germination, andthe attendant synthesis of proteins, than changes in abscisicacid content. Key words: Maize (Zea mays L.), germination, vivipary, desiccation, abscisic acid     

Seed storage conditions change the germination pattern of clonal growth plants in Mediterranean salt marshes

The effect of salinity level and extended exposure to different salinity and flooding conditions on germination patterns of three salt-marsh clonal growth plants (Juncus subulatus, Scirpus litoralis, and S. maritimus) was studied. Seed exposure to extended flooding and saline conditions significantly affected the outcome of the germination process in a different, though predictable, way for each species, after favorable conditions for germination were restored. Tolerance of the germination process was related to the average salinity level measured during the growth/germination season at sites where established individuals of each species dominated the species cover. No relationship was found between salinity tolerance of the germination process and seed response to extended exposure to flooding and salinity conditions. The salinity response was significantly related to the conditions prevailing in the habitats of the respective species during the unfavorable (nongrowth/nongermination) season. Our results indicate that changes in salinity and hydrology while seeds are dormant affect the outcome of the seed-bank response, even when conditions at germination are identical. Because these environmental-history-dependent responses differentially affect seed germination, seedling density, and probably sexual recruitment in the studied and related species, these influences should be considered for wetland restoration and management.