Seed dormancy and germination in three Crocus ser. Verni species (Iridaceae): implications for evolution of dormancy within the genus

The aim of this work was to examine whether seed ecophysiological traits in three closely related Crocus species were associated with ecological niche differentiation and species divergence. Seeds of the temperate tetraploid cytotype of Crocus neapolitanus, the sub‐Mediterranean C. etruscus and the Mediterranean C. ilvensis were placed either on agar in the laboratory under different periods of simulated seasonal conditions or in nylon mesh bags buried outdoors to examine embryo growth, radicle and shoot emergence. In agreement with the phenology observed outdoors, in the laboratory embryos required a cool temperature (ca. 10 °C) to grow to full size (embryo length:seed length, E:S ratio ca. 0.75) but only after seeds received a warm stratification; radicle emergence then followed immediately (November). Shoot emergence is a temporally separated phase (March) that was promoted by cold stratification in C. neapolitanus while in the other two species this time lag was attributed to a slow continuous developmental process. These species have similar embryo growth and radicle phenology but differ in their degree of epicotyl dormancy, which is related to the length of local winter. Conclusions from laboratory experiments that only consider root emergence could be misleading; evaluating the phenology of both root and shoot emergence should be considered in order to demonstrate ecologically meaningful differences in germination behaviour and to develop effective propagation protocols. Although these taxa resulted from recent speciation processes, the outcomes suggest an early onset of adaptation to local ecological factors and that phylogeny may represent a significant constraint in the evolution and expression of seed traits in Crocus.     

The Plight of Large Animals in Tropical Forests and the Consequences for Plant Regeneration

We introduce a special section that addresses the bushmeat or wild meat crisis, its direct impact on game species, and its indirect impact on plants in tropical forests.     

Effects of hydropriming treatments on the invigoration of aged Dodonaea viscosa seeds and water‐holding polymer on the improvement of seedling growth in a lava field

Restoration requires techniques similar to those used in agriculture to improve germination and seedling vigor. We treated 6‐year‐old (collected in 2003, S‐2003) and 1‐month‐old (S‐2009) seeds of Dodonaea viscosa with hydropriming (HP). Seeds were made permeable with hot water prior to hydration for 24 or 48 hours (HP‐24 and HP‐48, respectively) followed by dehydration. The resulting seedlings exposed to both HP treatments were sown in a lava field in soil mixed with hydrogel (HG) under the shade projected by five vegetation patches. The effects of these treatments on germination, seedling field survival, and growth were assessed. HP‐24 in S‐2009 and HP‐48 in S‐2003 increased the germination percentage from 22.5 and 31.7% in control seeds (permeable seeds) to 63.3 and 98.3%, respectively. The seedlings‐2009 (from S‐2009) with HG maintained high survival in all vegetation patches. Seedlings‐2003, however, had low survival. The lack of HG was negatively related to the photon flux in each patch. Survival of seedlings‐2009 increased with HG of up to 398.41 µmol m^?2 s^?1; after which survival decreased. During the rainy season, HP enhanced seedling growth, except the basal diameters and number of leaves in the seedlings‐2003 with HP‐24. During the dry season, the effects of HG and HP were similar for all the seedlings. In the following rainy season, the priming effect was lost while HG continued to promote seedling growth. The combined use of HP and HG and the shade projected by the patches resulted in a successful vegetation recovery strategy.     

Restoring montane cloud forest: establishment of three Fagaceae species in the old fields of central Veracruz,Mexico

Tropical montane cloud forest is rapidly disappearing and our knowledge of how to restore this system is limited. In a cloud forest of central Veracruz, Mexico, we studied seedling survival, growth, and causes of mortality (microenvironment and herbivory) of three native tree species, Fagus grandifolia, Quercus germana, and Q. xalapensis, transplanted into abandoned pastures (<1 year old) and secondary forests (9–17 years old). Microenvironment differed between the two habitats, temperature and photosynthetically active radiation were higher, and humidity was lower in the abandoned pastures than in the secondary forests. Seedling survival was greater in secondary forests than in pastures: F. grandifolia, 94 and 64%; Q. germana, 88 and 68%; and Q. xalapensis, 61 and 57%, respectively. The cause of mortality differed between habitats, with gophers (24.2% mortality) and mice (4.8%) killing the most seedlings in pastures, and damping‐off (16%) was the most important cause in secondary forests. The relative growth rate in height and basal area was significantly higher in abandoned pastures than in secondary forests; Q. xalapensis had the highest growth rate, followed by Q. germana and F. grandifolia. The environmental conditions in this mountainous cloud forest region seem less stressful to planted seedlings than the conditions of other lowland systems, as frequent clouds favor their establishment even in open sites. We conclude that Fagaceae species can successfully establish in abandoned pastures in mesic environments. Thus, the species studied can be used to speed cloud forest regeneration in the same area at different successional stages.     

Functions of canopy-stored seeds in the dune ecosystem: conclusions from Agriophyllum squarrosum and Artemisia wudanica

Delay in seed release is found in the dune ecosystem. However, its functions have been rarely reported. We studied delayed seed release in the Agrio-phyllum squarrosum (Chenopodiaceae), an annual psam-mophyte, and the Artemisia wudanica (Asteraceae), a psammophilous subshrub in an active sand dune field in Inner Mongolia, China, to get insights into its functions in dealing with sand movement. We concluded that delay in seed release in the dune ecosystem are as follows: (1) postponing dispersal of a portion of seeds until the end of windy season and the start of growing season; (2) regulating the spatiotemporal pattern of seed bank; (3) helping psammophytes adapt to wind erosion on the active sand dune; and (4) helping psammophytes realize long-distance dispersal.     

Environmental and genetic sources of diversification in the timing of seed germination: implications for the evolution of bet hedging

Environmental variation that is not predictably related to cues is expected to drive the evolution of bet-hedging strategies. The high variance observed in the timing of seed germination has led to it being the most cited diversification strategy in the theoretical bet-hedging literature. Despite this theoretical focus, virtually nothing is known about the mechanisms responsible for the generation of individual-level diversification. Here we report analyses of sources of variation in timing of germination within seasons, germination fraction over two generations and three sequential seasons, and the genetic correlation structure of these traits using almost 10,000 seeds from more than 100 genotypes of the monocarpic perennial Lobelia inflata. Microenvironmental analysis of time to germination suggests that extreme sensitivity to environmental gradients, or microplasticity, even within a homogeneous growth chamber, may act as an effective individual-level diversification mechanism and explains more than 30% of variance in time to germination. The heritability of within-season timing of germination was low (h(2) = 0.07) but significant under homogeneous conditions. Consistent with individual-level diversification, this low h(2) was attributable not to low additive genetic variance, but to an unusually high coefficient of residual variation in time to germination. Despite high power to detect additive genetic variance in within-season diversification, it was low and indistinguishable from zero. Restricted maximum likelihood detected significant genetic variation for germination fraction (h(2) = 0.18) under homogeneous conditions. Unexpectedly, this heritability was positive when measured within a generation by sibling analysis and negative when measured across generations by offspring-on-parent regression. The consistency of dormancy fraction over multiple delays, a major premise of Cohen's classic model, was supported by a strong genetic correlation (r = 0.468) observed for a cohort's germination fraction over two seasons. We discuss implications of the results for the evolution of bet hedging and highlight the need for further empirical study of the causal components of diversification.     

Agroecosystem management for rare species of Paysonia (Brassicaceae): integrating their seed ecology and life cycle with cropping regimens in a changing climate

Dormancy break and germination of seeds are governed by climatic cues, and predicted changes in climate may impact the ecology and conservation of species. Paysonia perforata and P. stonensis are rare brassicaceous winter annuals occurring primarily in fields on floodplains, where corn or soybeans are recommended for habitat maintenance. We tested the effects of precipitation, based on two predictions of changes in climate, on seed germination in these two species and placed the results into a management framework. Seeds of both species, collected during peak dispersal in late April/early May, were given various periods of light (or darkness) followed by darkness (or light) at summer temperatures before placement in darkness during late summer/early autumn in both laboratory and field. The light requirement was met earliest at 10 wk (mid-July) on alternating wet/dry substrate (simulating current climatic conditions). However, seeds of P. perforata and P. stonensis were photostimulated earliest at 2 wk (mid-May) and 6 wk (mid-June), respectively, on a continuously moist substrate (simulating predicted future conditions). The soil seed bank could be depleted if plowing coincides with photostimulation of seeds. Fields should be prepared after dispersal but before seeds are photostimulated and harvesting completed before seed germination in early September. Because seeds are highly photostimulated in late summer, disturbance from harvesting must be low to prevent burial. Cultivation of soybean, particularly for forage, is better matched to the seed biology and life cycle of Paysonia than that of corn under current and predicted climates.     

Testing the regeneration niche hypothesis with Gesneriaceae (tribe Sinningiae) in Brazil: Implications for the conservation of rare species

The germination requirements of sexually reproducing plants are regulated by environmental factors such as temperature. Those factors acting at the germination phase are part of the regeneration niche, which is fundamental in the processes that contribute to habitat suitability and geographic distribution. We tested the hypothesis that rarity is associated with regeneration niche in three species of plants in the family Gesneriaceae (tribe Sinningieae), Sinningia rupicola (Mart.) Wiehler, Paliavana sericiflora Benth and Sinningia allagophylla (Mart.) Wiehler, which vary in their distribution and habitat specificity but share a small zone of sympatry in rocky fields south of Belo Horizonte in Minas Gerais, Brazil. The regeneration niche was tested using a seed germination experiment under controlled light conditions at seven fixed temperatures (10–40°C at 5°C intervals). Each of the three species germinated differently at the various temperatures. The species with the smallest geographic range, S. rupicola, also had the most restricted germination: germination peaked at 15°C when relatively few seeds germinated (45%), and even fewer germinated at other temperatures. The regeneration niche was wider in P. sericiflora and wider still in S. allagophylla, with germination greater than 90% between 15–25°C and greater than 80% between 15–30°C, respectively. Our germination results provide qualified support for the hypothesis of correlation of the regeneration niche with geographic distribution of related plant taxa, with important conservation implications for rare and endangered species.     

A molecular framework for the embryo growth in germinating seeds of Solanum lycocarpum A. St.‐Hil., a nurse plant species

We are clarifying how the functional embryo growth occurs in germinating seeds of Solanum lycocarpum A. St.‐Hil., a nurse plant with a central role in forest dynamics in the Cerrado (a biodiversity hotspot). For that, we used classical seed germination measurements (germinability, mean germination time, mean germination rate, coefficient of variation of the germination time, synchronisation index and germination time range) and gene expression of mRNA codifying key proteins/enzymes for the success in the seed–seedling transition (Cyclin, Actin, Small Heat Shock Protein, Glutathione S‐transferase, Malate Dehydrogenase, Alcohol Dehydrogenase). Our findings demonstrate: (a) Although germination kinetics in S. lycocarpum seeds is slower than that in tomato seeds, the fold change of genes codifying key enzymes for the embryo development is similar in germinating seeds of both species. (b) The genes used here are useful, from a technical point of view, for classifying commercial seed samples of the species in relation to physiological quality. More notably, cyclin and malate dehydrogenase genes have a greater expression, both in germination sensu stricto and in immediate post‐germination. (c) A molecular framework for the embryo growth in germinating seeds of S. lycocarpum can be a functional explication for the species to be a nurse plant. Thus, the overlapping of classical and contemporary measurements is especially interesting to those species playing a central role in the environment, such as nurse plants, and may represent a new conservationist paradigm.     

The evolution of embryo size in angiosperms and other seed plants: implications for the evolution of seed dormancy

Abstract.— Seed dormancy plays an important role in germination ecology and seed plant evolution. Morphological seed dormancy is caused by an underdeveloped embryo that must mature prior to germination. It has been suggested that the presence of an underdeveloped embryo is plesiomorphic among seed plants and that parallel directional change in embryo morphology has occurred separately in gymnosperms and in angiosperms. We test these hypotheses using original data on embryo morphology of key basal taxa, a published dataset, and the generalized least squares (GLS) method of ancestral character state reconstruction. Reconstructions for embryo to seed ratio (E:S) using family means for 179 families showed that E:S has increased between the ancestral angiosperm and almost all extant angiosperm taxa. Species in the rosid clade have particularly large embryos relative to the angiosperm ancestor. Results for the gymnosperms show a similar but smaller increase. There were no statistically significant differences in E:S between basal taxa and any derived group due to extremely large standard errors produced by GLS models. However, differences between reconstructed values for the angiosperm ancestor and more highly nested nodes are large and these results are robust to topological and branch-length manipulations. Our analysis supports the idea that the underdeveloped embryo is primitive among seed plants and that there has been a directional change in E:S within both angiosperms and gymnosperms. Our analysis suggests that dormancy enforced by an underdeveloped embryo is plesiomorphic among angiosperms and that nondormancy and other dormancy types probably evolved within the angiosperms. The shift in E:S was likely a heterochronic change, and has important implications for the life history of seed plants.     

Seed germination in Chenopodium album L.: further evidence for the dependence of the effects of growth regulators on nitrate availability

Abstract Chenopodium album L. plants, grown under controlled environmental conditions on different levels of soil nitrate, produced seeds with proportionately different NO^?₃ contents. Regardless of the endogenous NO^?₃ content, few seeds germinated in water or upon treatment with KNO₃. Ethylene promoted germination, and the extent of germination was positively correlated with the endogenous seed NO^?₃ content. Combined application of ethylene and KNO₃ in the dark had a synergistic effect on NO^?₃ -deficient seed. The synergism between ethylene and KNO₃ was attributable to the NO^?₃ moiety of the nitrate salt. Ethylene and light showed moderate synergism in seeds with low or high endogenous nitrate. Addition of nitrate, however, masked the interaction between ethylene and light. Gibberellic acid₄₊₇ (GA₄₊₇) or red light, each alone or combined with KNO₃, had little effect on germination. When applied together in the dark, ethylene and GA₄₊₇ synergistically enhanced the germination of NO^?₃-deficient seed. The combined effects of the two hormones on this seed were further enhanced by the addition of KNO₃. There was no synergism between ethylene and GA₄₊₇ in NO^?₃-rich seed. These interactions among GA₄₊₇, ethylene and KNO₃ were not affected by light. The results confirm and further elaborate our earlier finding that the sensitivity of C. album seeds to ethylene may depend on nitrate availability.     

The Shape of the Female Fitness Curve for Cynoglossum officinale: Quantifying Seed Dispersal and Seedling Survival in the Field

Abstract: When hermaphroditic plants shift their sex allocation to produce more seeds and less pollen, it is frequently assumed that the female component of fitness is strictly proportional to the number of seeds produced. However, if producing more seeds results in more competition between seedlings, the female fitness gain curve levels off with high investments in seeds. The shape of this curve is relevant for sex allocation theory, but rarely have data been collected. For Cynoglossum officinale we described the relationship between the number of seeds produced on the mother plant and the number and weight of seedlings in September of the following year. As expected, around large plants of C. officinale more seedlings were retrieved after the germination period in March. The seedlings of large plants were dispersed over an area similar to that of small plants. As a result, seedlings around large plants had a significantly higher chance to have a neighbouring sibling within one dm². Survival and growth of single or grouped (density > 2) seedlings did not differ significantly. As a result, total dry weight of seedlings in September was a linear function of the number of seeds on the parent plant. Our data indicate a linear female fitness gain curve.     

Effects of climate warming on plant autotoxicity in forest evolution: a case simulation analysis for Picea schrenkiana regeneration

In order to explore how plant autotoxicity changes with climate warming, the autotoxicity of P. schrenkiana needles' water extract, organic extract fractions, and key allelochemical DHAP was systemically investigated at the temperature rising 2 and 4°C based on the data‐monitored soil temperature during the last decade in the stage of Schrenk spruce regeneration (seed germination and seedling growth). The results showed that the criterion day and night temperatures were 12°C and 4°C for seed germination, and 14°C and 6°C for seedling growth, respectively. In the presence of water extract, the temperature rise of 2°C significantly inhibited the germination vigor and rate of P. Schrenkiana seed, and a temperature rise of 4°C significantly increased the inhibition to the seedling growth (P < 0.05). Among the three organic fractions, the low‐polar fraction showed to be more phytotoxic than the other two fractions, causing significant inhibitory effects on the seed germination and growth even at low concentration of 0.1 mg/mL, and the inhibition effect was enhanced as temperature increased. The temperature rise significantly enhanced the promotion effect of DHAP, while the inhibition effect of temperature rise became less important with increasing concentration of DHAP. This investigation revealed that autotoxicity of P. schrenkiana was affected by the climate warming. As expected, it provided an insight into the mechanism and effectiveness of allelopathy in bridging the causal relationship between forest evolution and climate warming.     

Genetic diversity of globe artichoke landraces from Sicilian small-holdings: implications for evolution and domestication of the species

Globe artichoke (Cynara cardunculus var. scolymus) is native to the Mediterranean Basin, where it grows in close proximity with its ancestor wild cardoon (C. cardunculus var. sylvestris); its commercial production is mainly based on vegetatively propagated clones which guarantee high yields of marketable product (i.e. immature inflorescence or capitula). A collection of 24 landraces of globe artichoke was made from small-holdings in Sicily, which is assumed to be one of the possible centres of its domestication. These landraces have been cultivated for centuries by local farmers, mainly due to their culinary uniqueness. The collection was characterised for a combination of morphological traits and AFLP, gSSR and cpSSr markers. Molecular analyses included genotypes of wild cardoon collected from different sites in Sicily as well as accessions of the most widely grown Sicilian varietal types: the spiny ‘Spinoso di Palermo’ and the non-spiny ‘Violetto di Sicilia’. The landraces follow a gradient of ‘ennoblement’ towards either the domesticated spiny or the non-spiny types. ‘Cimiciusa di Mazzarino’ was an outlier, in that it resembled the cultivated forms with respect to its AFLP fingerprint, but was more closely related to the wild cardoon on the basis of SSR profile. This particular landrace presents an example of an intermediary form in the domestication process, although it could also have derived from introgression from sympatric wild cardoon, followed by farmer selection. The abundant genetic variation present demonstrates the key role of farmers’ practice in the maintenance of genetic diversity, which should be preserved because of its potential value for plant breeders.     

Molecular evolution of the hyaluronan synthase 2 gene in mammals: implications for adaptations to the subterranean niche and cancer resistance

The naked mole-rat (NMR) Heterocephalus glaber is a unique and fascinating mammal exhibiting many unusual adaptations to a subterranean lifestyle. The recent discovery of their resistance to cancer and exceptional longevity has opened up new and important avenues of research. Part of this resistance to cancer has been attributed to the fact that NMRs produce a modified form of hyaluronan—a key constituent of the extracellular matrix—that is thought to confer increased elasticity of the skin as an adaptation for living in narrow tunnels. This so-called high molecular mass hyaluronan (HMM-HA) stems from two apparently unique substitutions in the hyaluronan synthase 2 enzyme (HAS2). To test whether other subterranean mammals with similar selection pressures also show molecular adaptation in their HAS2 gene, we sequenced the HAS2 gene for 11 subterranean mammals and closely related species, and combined these with data from 57 other mammals. Comparative screening revealed that one of the two putatively important HAS2 substitutions in the NMR predicted to have a significant effect on hyaluronan synthase function was uniquely shared by all African mole-rats. Interestingly, we also identified multiple other amino acid substitutions in key domains of the HAS2 molecule, although the biological consequences of these for hyaluronan synthesis remain to be determined. Despite these results, we found evidence of strong purifying selection acting on the HAS2 gene across all mammals, and the NMR remains unique in its particular HAS2 sequence. Our results indicate that more work is needed to determine whether the apparent cancer resistance seen in NMR is shared by other members of the African mole-rat clade.     

Postdispersal seed predation and seed viability in forest soils: implications for the regeneration of tree species in Ethiopian church forests

Almost all dry Afromontane forests of Northern Ethiopia have been converted to agricultural, grazing or scrub lands except for small fragments left around churches (‘Church forests’). Species regeneration in these forests is limited. We investigated (i) how intense postdispersal seed predation was in church forest, and if this seed predation varied with species and/or habitat, and (ii) for how long tree seeds maintained their viability while buried in forest soil. In the seed predation experiment, we monitored seeds of six tree species in four habitats for a period of 14 weeks (the peak seeding season). In the seed viability experiment, we assessed seed viability of five species in four habitats after being buried 6, 12, or 18 months. Ninety‐two percent of the tree seeds were predated within 3.5 months. Predation was mainly dependent on species whereas habitat had a weaker effect. Seed viability decreased sharply with burial time in soil for all species except for Juniperus. To minimize seed availability limitation for regeneration of such species in the forest, the standing vegetation needs to be persistently managed and conserved for a continuous seed rain supply. Additional seed sowing, and seed and seedling protection (by e.g. animal exclosures) may increase successful regeneration of important species in these forests.     

Morphological structure of the aortic wall in three Delphinid species with shallow or intermediate diving habits: Evidence for diving adaptation

Some modifications in the vascular system of marine mammals provide adaptive advantages for diving. This study analyses the organisation of the aortic wall in dolphins, observing artery changes in volume and blood pressure for diving behaviour. Samples of three aortic segments (ascending, thoracic and abdominal) of three dolphin species were processed for histological and morphometric studies. The three dolphin species used, striped dolphin (Stenella coeruleoalba), Atlantic spotted dolphin (Stenella frontalis) and common dolphin (Delphinus delphis), have shallow or intermediate diving habits. Our results indicated that the components of the aortic wall of the dolphins had different dispositions in the three selected segments. The aortic wall decreased in thickness along its length due to a loss of the lamellar units in the tunica media and a thinning of the main elements of the lamellar units along the artery. The life stage had little influence on the thickness of the aortic wall except for the ascending aorta. The weight, body length, species or sex of the specimen did not significantly influence the thickness of the wall or the lamellar units. In summary, the histological and morphometric aortic structure in dolphins, in relation to the studied parameters, seems to be similar to that previously described of terrestrial mammals such as pigs, except for a larger difference in the proportion of lamellar units between the ascending and thoracic segments.