At the core of plant regeneration, temperature and water supply are critical drivers for seed dormancy (initiation, break) and germination. Hence, global climate change is altering these environmental cues and will preclude, delay, or enhance regeneration from seeds, as already documented in some cases. Along with compromised seedling emergence and vigour, shifts in germination phenology will influence population dynamics, and thus, species composition and diversity of communities. Altered seed maturation (including consequences for dispersal) and seed mass will have ramifications on life history traits of plants. Predicted changes in temperature and precipitation, and thus in soil moisture, will affect many components of seed persistence in soil, e.g. seed longevity, dormancy release and germination, and soil pathogen activity. More/less equitable climate will alter geographic distribution for species, but restricted migratory capacity in some will greatly limit their response. Seed traits for weedy species could evolve relatively quickly to keep pace with climate change enhancing their negative environmental and economic impact. Thus, increased research in understudied ecosystems, on key issues related to seed ecology, and on evolution of seed traits in nonweedy species is needed to more fully comprehend and plan for plant responses to global warming.
The most often used time-line for distinguishing a transient seed bank from a persistent seed bank is one calendar year. Thus, species whose seeds live in or on the soil for <1 year have a transient seed bank, whereas those whose seeds live for ≥1 year have a persistent seed bank. However, dormancy cycling of seeds buried in soil has not been given due consideration in these models. When dormancy cycling is considered, it is shown that seeds of both autumn-germinators and spring-germinators are in the dormant state when they are 1 year old. Thus, unless the seeds live until at least the second germination season (i.e. usually 16–18 months following dispersal), they are, in effect, part of a transient seed bank, having lived through only one germination season. We propose that for seeds of such species to be considered part of a short-term persistent seed bank, they should remain viable and germinable until at least the second germination season, and to be part of a long-term persistent seed bank, until at least the sixth germination season. Our definitions are applicable to seeds with physiological, physical or morphophysiological dormancy, which often require >1 year after maturity to come out of dormancy in nature. We discuss modifications of the seedling emergence method for detection of a soil seed bank, so that they correspond to our definitions of seed-bank strategies.
Osmorhiza aristata is an herbaceous perennial that grows primarily in Japan, through southern China, to the Himalayas. It closely resembles the eastern North American species O. claytonii and O. longistylis, and, together, the three species are an example of the well-known North American-Asian pattern of disjunction. Requirements for dormancy break and embryo growth were determined for seeds of O. aristata collected in Japan during the summers of 1998-2000. Embryos in fresh seeds were ca. 0.5 mm long, and they had to grow to 9 mm before the radicle emerged from the mericarp. Embryo growth and germination occurred during cold stratification at 5°C, the optimum temperature for germination. Gibberellic acid did not substitute for cold stratification. Thus, O. aristata seeds have deep complex morphophysiological dormancy (MPD). The type of MPD in O. aristata is similar to that in two western North American congeners but different from that in eastern North American congeners (nondeep complex MPD). Mapping the types of MPD onto a phylogeny of the genus suggests that nondeep complex MPD is derived from deep complex MPD. Although eastern North American-Asian disjuncts often exhibit morphological stasis, the taxa may differ greatly in physiological traits, such as seed dormancy.
Although the study species are con-generic, sympatric and produce seeds of identical morphology, they possessed different dormancy-break and germination requirements. The physiological component of MPD was non-deep in H. racemosa but varied in the other three species where more deeply dormant seeds required >1 summer to overcome dormancy and, thus, germination was spread over time. Embryos grew during winter, but future studies need to resolve the role of cold versus warm stratification by using constant temperature regimes. To include Mediterranean species with MPD, some modifications to the current seed-dormancy classification system may need consideration: (a) wet/dry conditions for warm stratification and (b) a relatively long period for warm stratification. These outcomes have important implications for improving experimental approaches to resolve the effective use of broadcast seed for ecological restoration.
Dormancy-breaking requirements and types of dormancy were determined for seeds ofLonicera fragrantissimaLindl. & Paxt.,L. japonicaThunb.,L. maackii(Rupr.) Maxim. andL. morrowiiA. Gray. Seeds of all four species have underdeveloped spatulate embryos that are about 20–40%fully developed (elongated) when dispersed. Embryos in freshly matured, intact seeds grew better at 25/15°C than at 5°C. Gibberellic acid (GA3) (tested only in the light) was more effective in breaking dormancy inL. maackiiandL. morrowiithan inL. fragrantissimaandL. japonica. Warm- followed by cold stratification was required to break dormancy in seeds ofL. fragrantissima, whereas seeds ofL. japonicarequired cold stratification only. Thus, seeds ofL. fragrantissimahave deep simple morphophysiological dormancy (MPD) and those ofL. japonicanondeep simple MPD. About 50%of the seeds ofL. maackiirequired warm- or cold stratification only to come out of dormancy and 50% of those ofL. morrowiirequired warm stratification only, whereas the other 50% did not require stratification to germinate. Thus, about half of the seeds of the two species has nondeep simple MPD, and the other half has morphological dormancy (MD). In these laboratory tests, seeds ofL. japonica,L. maackii, andL. morrowiigenerally germinated to significantly higher percentages in light than in darkness; seeds ofL. fragrantissimawere not tested in darkness. Peaks of germination for seeds ofL. fragrantissima,L. japonica,L. maackiiandL. morrowiisown on a soil surface and covered withQuercusleaves under near-natural temperature conditions shortly after seed maturity and dispersal in late June 1997, late November 1997, early November 1996 and late June 1998, respectively, occurred in early March 1998, late February 1998, late March 1997 and early October 1998, respectively. The germination phenologies of seeds of the same species and seed lots buried in soil were similar to those of seeds under leaf litter. High percentages of seeds of all four species germinated both under litter (78–96%) and beneath the soil surface (78–97%). These germination patterns correspond closely with the requirements for embryo growth and dormancy break in the fourLoniceraspecies.
Requirements for dormancy break and embryo growth were determined for seeds of the western North American species, Osmorhiza depauperata. Seeds were collected in August 2001 from Sandia Crest (3200 m elevation) and Las Huertas (2300 m), New Mexico (USA). Embryos in fresh seeds were c. 0.6 mm long, and they had to grow to c. 9–10 mm before the radicle emerged from the mericarp. Embryo growth occurred at low temperatures (1 and 5°C), and seeds germinated to high percentages at 1°C during 32 weeks of incubation in the light. No seeds germinated at 5, 15/6, 20/10, 25/15 or 30/15°C during 32 weeks of incubation. Although a 4–18 week warm-temperature (25/15°C) pretreatment increased germination rates at 1°C, it was unnecessary for a high percentage of seeds to germinate. Gibberellic acid (GA3, 10–1000 mg l–1) did not substitute for cold stratification. Seeds from the low-elevation population contained larger embryos and required less time to germinate than those from the high-elevation population. O. depauperata seeds have deep complex morphophysiological dormancy (MPD), which is similar to two other western North American congeners and an Asian congener, but different from two eastern North American congeners. Results from this study suggest that: (1) phylogenetic niche conservatism has played a role in the persistence of deep complex MPD in the three western North American species of Osmorhiza; and (2) the stimulatory effect from a warm pretreatment in species needing only cold stratification for dormancy break is a preadaptation that initiated the development of an absolute warm requirement in species needing both warm and cold stratification.
In contrast to previous reports, the endocarps ("seed coats") of Sambucus species are not impermeable to water; thus, the seeds do not have physical dormancy. Seeds of the North American species Sambucus canadensis and S. pubens and of the European species S. racemosa have spatulate shaped embryos that are ∼60% fully developed (elongated) at seed maturity. The embryo has to extend to the full length of the seed to germinate. Embryos in freshly matured seeds of S. canadensis and in those of S. pubens grew better at 25°/15°C than at 5°C, whereas the rate of embryo growth in S. racemosa was higher at 5°C than at 25°/15°C. Seeds of all three species germinated to significantly higher percentages in light (14-h photoperiod) than in darkness. Fresh seeds of neither species germinated during 2 wk of incubation over a range of thermoperiods. Warm followed by cold stratification broke dormancy in seeds of S. canadensis and in those of S. pubens. Thus, seeds of these two North American species have deep simple morphophysiological dormancy (MPD). In comparison, seeds of the European species S. racemosa required a cold stratification period only for dormancy break, and thus they have intermediate complex MPD. GA(3) was much more effective in breaking dormancy in seeds of S. racemosa than it was in those of S. canadensis or S. pubens.
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