Physical dormancy (PY) in seeds/fruits, which is caused by the water-impermeable palisade layer, has long been considered a mechanism for synchronizing germination to a favourable time for seedling survival and establishment. Recently, a new hypothesis (crypsis hypothesis) was proposed as the main selective factor for the evolution of PY. However, there are some misconceptions in this hypothesis. Our objective is to critically evaluate the crypsis hypothesis and to point out that there are multiple adaptive roles of PY. The fundamental argument in the crypsis hypothesis, that PY evolved as an escape mechanism from predators, is not valid according to the evolutionary theory of Darwin. According to Darwin's hypothesis, variations (dormancy in our case) within a population occur randomly, i.e. there is no direct function of a variation at the time of its origin. Different selection pressures operating in the environment increase or decrease the fitness of individuals with the variation. Water-gap anatomy in seeds/fruits and phylogenetic relationships of species with PY suggest that PY has evolved several times in angiosperms. Thus, we argue that not only predatory pressure but also several other environmental pressures (e.g. proper timing of germination, ultra-drying of seeds, dispersal and pathogens) were involved in increasing the fitness of species producing seeds with PY. The significance of PY in the survival of the species under the abovementioned environmental pressures and other misconceptions of the crypsis hypothesis are discussed in detail.
Community‐level seed dormancy studies are important in understanding the dynamics of plant communities and adaptations of species to their habitat. Our aim was to develop a seed dormancy profile for tropical montane forests of Sri Lanka, which are a global biodiversity hotspot, and compare it to the profile made using a world database for this vegetation type. Germination, imbibition and embryo length : seed length ratio of seeds were determined for 80 Sri Lankan montane forest species. Seeds of 31 species were fast‐germinating with a median length of germination (MLG) <30 days and the remaining 49 were slow‐germinating with MLG >30 days. Embryos of six fast‐germinating species grew prior to radicle emergence, indicating morphological dormancy (MD). The other 25 fast‐germinating species had non‐dormant (ND) seeds. Manually scarified seeds of two species imbibed significantly more water than non‐scarified seeds, revealing physical dormancy (PY). Embryos of 20 slow‐germinating species grew prior to radicle emergence, confirming morpho‐physiological dormancy (MPD). The remaining slow‐germinating species had a fully developed embryo and thus physiological dormancy (PD). The percentage of species with ND seeds and with MD, MDP, PD and PY was 31, 7.5, 25, 34 and 2.5, respectively. Species with dormant seeds (70%) dominate the Sri Lankan montane forest community similar to the world database, with 85% dormant seeds. Seed dormancy may be an adaptation that prevents seeds from germinating during the Sri Lankan dry season from December to March when conditions are unfavourable for seedling growth due to low water availability.
To increase our knowledge of the diversity of seed dormancy and germination in Rubiaceae, we investigated seed desiccation sensitivity and germination of threePsychotriaspecies. Seeds ofP. gardneri, P. nigraandP. zeylanicagerminated to high percentages at <15% seed moisture content. Intact seeds ofP. zeylanicaandP. nigraimbibed water and thus do not have physical dormancy. More than 50% of the seeds ofP. zeylanica, P. nigraandP. gardneritook 33, 53 and 110 d, respectively, at 25°C for the radicle to emerge, and embryo growth occurred before and after radicle emergence. Thus, seeds have morphophysiological dormancy. Shoot emergence ofP. nigraandP. zeylanicaseeds was delayed 50 and 80 d after radical emergence, respectively; thus, seeds have epicotyl morphophysiological dormancy (eMPD). This is the first report of eMPD in Rubiaceae. Since warm stratification promoted both radicle and shoot emergence in seeds ofP. zeylanicaandP. nigra, the level of eMPD is non-deep simple. Hence, dormancy of the studiedPsychotriaspp. can be described as C1bBb(radicle)–C1bBb(epicotyl), i.e. the embryo is underdeveloped and grows prior to radicle emergence and epicotyl emergence under warm temperatures (Bb), and both the radicle and epicotyl have non-deep simple physiological dormancy broken by warm temperatures (C1b). In twoPsychotriaspecies studied in detail, radicle emergence occurs at the beginning of the rainy season and plumule emergence at the peak rainy season when conditions are most favourable for rapid seedling development.
Fruiting season of many Sri Lankan tropical montane species is not synchronised and may not occur when conditions are favourable for seedling establishment. We hypothesised that species with different fruiting seasons have different seed dormancy mechanisms to synchronise timing of germination with a favourable season for establishment. Using six species with different fruiting seasons, we tested this hypothesis. Germination and imbibition of intact and manually scarified seeds were studied. Effect of GA on germination was examined. Embryo length:seed length (E:S) ratio of freshly matured seeds and of those with a split seed coat was determined. Time taken for radicle and plumule emergence and morphological changes of the embryos were recorded. The radicle emerged from Ardisia missionis, Bheza nitidissima and Gaetnera walkeri seeds within 30 days, whereas it took >30 days in other species. Embryos grew in seeds of B. nitidissima and G. walkeri prior to radicle emergence but not in Microtropis wallichiana, Nothapodytes nimmoniana and Symplocos cochinchinensis. A considerable delay was observed between radicle and plumule emergence in all six species. Warm stratification and/or GA promoted germination of all species. All the tested species have epicotyl dormancy. Seeds of B. nitidissima and G. walkeri have non-deep simple morphophysiological epicotyl dormancy, and the other four species have non-deep physiological epicotyl dormancy. Differences in radicle and epicotyl dormancy promote synchronisation of germination to a favourable time for seedling development. Therefore, information on dormancy-breaking and germination requirements of both radicle and epicotyl are needed to determine the kind of dormancy of a particular species.
Although the level of seed desiccation sensitivity (LSDS) may have an impact on plant species conservation, information is available for <10% of tropical angiosperms. A study was conducted to assess the LSDS of 28 tropical montane species in Sri Lanka. Seeds were extracted from freshly collected fruits. Initial weight was recorded, and thousand seed weight (TSW) was calculated. Seed moisture content (MC) was determined. LSDS was determined using seed desiccation experiments and predicted using the TSW–MC criterion. Seed storage behaviour was predicted using LSDS and storage data and using a model based on phylogenetic affiliation. The relationship between LSDS and seed dormancy, life form and forest strata was evaluated. Fresh seeds of only 12 species germinated to >80%. Although seeds of the other species had >80% viability, only 0–70% germinated due to dormancy. Seeds of five species had MC <15%, indicating desiccation tolerance (DT). Seeds of 12 species lost viability after desiccation, indicating desiccation sensitivity (DS). Seeds of Ardisia missionis, Psychotria gartneri and Psychotria nigra remained viable after desiccation, showing DT. Seeds of 17 species were DS and those of 11 species DT. The TSW of four species was >500 g. Thus, seeds of other species were predicted to be DT by the TSW–MC criterion. A relationship was identified between LSDS and the forest strata of the species. More canopy species produced DS than DT seeds. Since seeds of most of the studied species were DS, these species may be threatened due to prolonged droughts predicted for the region due to climate change.
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