Seed Germination Ecology of the Cold Desert Annual Isatis violascens (Brassicaceae): Two Levels of Physiological Dormancy and Role of the Pericarp

The occurrence of various species of Brassicaceae with indehiscent fruits in the cold deserts of NW China suggests that there are adaptive advantages of this trait. We hypothesized that the pericarp of the single-seeded silicles of Isatis violascens restricts embryo expansion and thus prevents germination for 1 or more years. Thus, our aim was to investigate the role of the pericarp in seed dormancy and germination of this species. The effects of afterripening, treatment with gibberellic acid (GA₃) and cold stratification on seed dormancy-break were tested using intact silicles and isolated seeds, and germination phenology was monitored in an experimental garden. The pericarp has a role in mechanically inhibiting germination of fresh seeds and promotes germination of nondormant seeds, but it does not facilitate formation of a persistent seed bank. Seeds in silicles in watered soil began to germinate earlier in autumn and germinated to higher percentages than isolated seeds. Sixty-two percent of seeds in the buried silicles germinated by the end of the first spring, and only 3% remained nongerminated and viable. Twenty to twenty-five percent of the seeds have nondeep physiological dormancy (PD) and 75–80% intermediate PD. Seeds with nondeep PD afterripen in summer and germinate inside the silicles in autumn if the soil is moist. Afterripening during summer significantly decreased the amount of cold stratification required to break intermediate PD. The presence of both nondeep and intermediate PD in the seed cohort may be a bet-hedging strategy.     

Seed germination during floatation and seedling growth of Carapa guianensis, a tree from flood-prone forests of the Amazon

Carapa guianensis Aubl. (Meliaceae), a hard wood tree from the Brazilian Amazon, has large recalcitrant seeds that can germinate and establish in both flood-free (terra-firme) and flood-prone (várzea) forests. These seeds, although large, can float. This study was designed to experimentally examine seed longevity under floating conditions ex-situ and its effects on subsequent germination and seedling growth. Many seeds germinated while floating, and radicle protrusion occurred from 3 to 42 d after the start of the floating treatment (tap water, room temperature 20–30 °C). Shoots of newly germinated floating seedlings may elongate up to 37.0 cm in 20 d without loss of viability. Epicotyl and first leaf emergence were delayed by floating. Seeds that did not germinate while floating were then placed on vermiculite and watered daily, where many seeds resumed germination. Germination during and after floating was affected by the length of the floating treatment: 88% germinated after 1 mo, 82% germinated after 2 mo and 70% germinated after 2.5 mo. These results indicate that Carapa guianensis has physiological variation regarding dormancy in response to seed floatation. The fact that floatation induces dormancy in recalcitrant seeds of this economically important species can be relevant to initiatives of ex situ storage of seeds.     

Dormancy-breaking and germination requirements for seeds of Symphoricarpos orbiculatus (Caprifoliaceae)

Fruits (drupes) of Symphoricarpos orbiculatus ripen in autumn and are dispersed from autumn to spring. Seeds (true seed plus fibrous endocarp) are dormant at maturity, and they have a small, linear embryo that is underdeveloped. In contrast to previous reports, the endocarp and seed coat of S. orbiculatus are permeable to water; thus, seeds do not have physical dormancy. No fresh seeds germinated during 2 wk of incubation over a 15°/6°-35°/20°C range of thermoperiods in light (14-h photoperiod); gibberellic acid and warm or cold stratification alone did not overcome dormancy. One hundred percent of the seeds incubated in a simulated summer → autumn → winter → spring sequence of temperature regimes germinated, whereas none of those subjected to a winter → spring sequence did so. That is, cold stratification is effective in breaking dormancy only after seeds first are exposed to a period of warm temperatures. Likewise, embryos grew at cold temperatures only after seeds were exposed to warm temperatures. Thus, the seeds of S. orbiculatus have nondeep complex morphophysiological dormancy. As a result of dispersal phenology and dormancy-breaking requirements, in nature most seeds that germinate do so the second spring following maturity; a low to moderate percentage of the seeds may germinate the third spring. Seeds can germinate to high percentages under Quercus leaf litter and while buried in soil; they have little or no potential to form a long-lived soil seed bank.     

Germination of Prosopis juliflora (Sw) DC seeds after scarification treatments

Invasive plant species are the second most important threat to global biodiversity loss after land‐use change. Invasive species can modify native community composition, deplete species diversity and affect ecosystem processes. The Caatinga is one of the most human‐affected Brazilian ecosystems owing to non‐sustainable use of its natural resources. Prosopis juliflora is an important invasive plant species in the Caatinga ecosystem. Seed germination is a critical stage in plant life cycles and is a major factor in the establishment and success of invasive plant species. Among the factors that affect seed germination and dormancy, coat‐imposed seems to be the most important for P. juliflora. In Prosopis species, the ingestion of fruits by wild and domestic animals may promote and accelerate germination, enhancing the dispersal of seeds and fruits of these species. We investigated the germination capacity of P. juliflora seeds after artificial mechanical and chemical scarification and analyzed the changes in seedling vigor caused by the scarification treatments. Germination rate, germination time (TMG) and germination synchrony (E) differed significantly with the length of the scarification treatments in H₂SO₄ for both seeds with endocarps and seeds without endocarps (non‐endocarp seeds). Sulfuric acid affected plant survival more strongly than germination rate, particularly in non‐endocarp seeds.     

Role of temperature in the germination ecology of three summer annual weeds

Summary Ambrosia artemisiifolia L., Chenopodium album L., and Amaranthus retroflexus L. are three summer annual weeds that occur in disturbed habitats. In nature, the peak germination season for A. artemisiifolia and C. album is in early to mid-spring, while in A. retroflexus the peak germination season is late spring to early summer. Furthermore, seeds of A. artemisiifolia germinate only in spring, while seeds of C. album and A. retroflexus germinate throughout the summer. In an attempt to explain the differential germination behavior of these three species in nature, changes in their germination responses to temperature during burial in a non-heated greenhouse from October 1974 to October 1975 were monitored. A high percentage of the seeds of all three species after-ripened during winter. Seeds of A. artemisiifolia and C. album germinated at temperatures characteristic of those in the field in early and mid-spring, but seeds of A. retroflexus required the higher temperatures of late spring and early summer for germination. Seeds of all three species germinated to higher percentages in light than in darkness. Non-dormant seeds of A. artemisiifolia that did not germinate in spring entered secondary dormancy. On the other hand, seeds of C. album and A. retroflexus that did not germinate when temperatures first became favorable for germination, did not enter secondary dormancy and, thus, retained the ability to germinate at summer field temperatures during summer. Thus, temporal differences in the germination behavior of these three species are caused by the differential reaction of the seeds to temperature during the annual temperature cycle.     

Ecophysiology of seed germination in Erythronium japonicum (Liliaceae) with underdeveloped embryos

Erythronium japonicum (Liliaceae) (Japanese name, katakuri) is indigenous to Japan and adjacent Far East regions. We examined their embryo elongation, germination, and seedling emergence in relationship to the temperature. In incubators, seeds did not germinate at 20°/10° (light 12 h/dark 12 h alternating temperature), 20°, 15°, 5°, or 0°C with a 12-h light photoperiod for 200 d. They germinated at 15°/5° or 10°C, starting on day 135. If seeds were kept at 20° or at 25°/15°C before being exposed to 5°C, the seeds germinated, but if kept at 25° or 30°C they did not. Embryos at 25°/15°C grew to half the seed length without germinating; at 0° or 5°C, embryos elongated little. Embryos grew and seeds germinated when kept at 25°/15°C for 90 d and then at 5°C. In the field, seeds are dispersed in mid-June in Hokkaido and in Honshu, mid-May to mid-June. Seeds do not germinate immediately after dispersal because the embryo is underdeveloped. Embryos elongated at medium temperatures in autumn after summer heat, and germination ends in November at 8°/0°C. After germination, seedling emergence was delayed, and most seedlings were observed in early April around the snowmelt when soil cover was 2-3 mm.     

DORMANCY AND GERMINATION IN SEEDS OF COMMON RAGWEED WITH REFERENCE TO BEAL'S BURIED SEED EXPERIMENT

Common ragweed (Ambrosia artemisiifolia L.) was one of 19 herbaceous weedy species used by Beal in his buried viable seed experiment started in 1879. No seeds germinated during the first 35 years of the experiment when germination tests were performed in late spring, summer or early autumn. Germination did occur in seeds buried for 40 years when seeds were exhumed and tested for germination in early spring. Data obtained in more recent research provide the probable explanation for these results. Seeds of common ragweed that do not germinate in spring enter secondary dormancy by mid to late spring and will not germinate until dormancy is broken the following late autumn and winter. Thus, during the first 35 years of the experiment seeds were dormant when tested for germination, whereas seeds buried for 40 years were nondormant. Seeds buried 50 years or longer did not germinate when tested in spring, probably because they had lost viability and/or seeds germinated during burial and seedlings died.     

GERMINATION ECOPHYSIOLOGY OF THE WOODLAND HERB OSMORHIZA LONGISTYLIS (UMBELLIFERAE)

Osmorhiza longistylis is an herbaceous perennial that grows in woodlands of eastern and central North America. In northcentral Kentucky seeds ripen in early to mid July, and dispersal begins in September and October. Although most of the seeds are shed during late autumn and winter, some remain on the dead shoots for up to 18 months. Seeds are dormant at maturity due to an underdeveloped embryo. Embryos grew at low (5 C) temperatures, but only after seeds were given a period of warm (30/15 C) stratification. With an increase in the length of the warm treatment, there was an increase in the number of embryos that grew to full length during a 12-wk period at 5 C and an increase in the percentage of seeds that germinated. Seeds given 12 wk of warm stratification required more than 8 wk at 5 C to overcome dormancy. Embryos in freshly-matured seeds averaged 0.60 mm long, but those in seeds given 12 wk warm plus 12 wk cold stratification averaged 8.86 mm. Lengths of embryos of seeds kept moist at 30/15 and 5 C for 24 wk averaged 0.63 and 0.89 mm, respectively. Regardless of age and dispersal time, imbibed seeds must be exposed to high (i.e., summer or autumn) and then to low (i.e., winter) temperatures before they will germinate. Consequently, germination occurs only in spring.     

Sterile Germination Requirements of Seeds of Some Water Plants

A sterile germination study with seeds of some common phanerogamic water plants showed almost 100 per cent germination for seeds of Alisma plantago–aquatica, Baldellia ranunculoides and Nymphaea alba. Seeds of Potamogeton lucens could be germinated to about 40 per cent, seeds of Polygonum amphibium germinated sporadically while those of Cladium mariscus could not be germinated at all. Freshly harvested seeds of Alisma and Baldellia showed an ability to germinate at both 20°C and 35°C. A stratification period of one month at +4°C gave germination of all species tested, with the exception of Cladium. Potamogeton germinated in light only, the other species both in light and darkness. Treatment times for surface sterilization in disinfectants are given.     

Seed production,dispersal and germination in Cryptostegia grandiflora and Ziziphus mauritiana,two invasive shrubs in tropical woodlands of northern Australia

Abstract Cryptostegia grandiflora and Ziziphus mauritiana are exotic shrubs that are invading tropical woodlands of northern Australia. Although they are widespread, they currently occupy only small proportions of their respective potential ranges. Large C. grandiflora can produce more than 8000 wind-dispersed seeds in a single reproductive episode and can set seed at least twice per year. More than 90% of seeds will germinate within 10 days of moisture becoming available. Few, if any, seeds survive for more than 12 months in the soil. Large Z. mauritiana can produce more than 5000 fruits per year. The fruit is a drupe that contains a single seed (sometimes two) enclosed in a woody endocarp. Less than 10% of fresh seeds will germinate without removal of the endocarp. Removal of the endocarp increased germination to 56%. Less than 10% of seeds of Z. mauritiana remain viable after burial in the soil for 12 months. Seeds of Z. mauritiana are dispersed by several mammalian vectors, including wallabies and feral pigs, but domestic cattle are probably the major means by which large numbers of seeds reach new sites. Propagules pass intact through the digestive tract of cattle, feral pigs and wallabies and contain viable seeds that germinate more readily than seed in fresh intact fruits. Controlling movement of cattle that have had recent access to mature fruit of Z. mauritiana could significantly reduce the likelihood of new infestations. For both species, management that reduces seed production will be important for containing spread.     

Breaking Setaria parviflora seed dormancy by nitrates and light is part of a mechanism that detects a drawdown period after flooding

We explored the hypothesis that, in flood-prone habitats, nitrates can signal to seeds that a drawdown period has begun. To investigate this issue, Setaria parviflora (Poir.) Kerguélen seeds were buried in a never-flooded upland and a nearby, flood-prone lowland grassland. Seeds were exhumed during the flooding period. Additionally, grassland mesocosms with buried S. parviflora seeds were flooded during 20 d (controls drained). After both field and mesocosm pretreatments germination was assayed in laboratory at 25 °C in a medium with or without nitrates, under red light pulses or in darkness. Seeds exhumed from the never-flooded upland showed no specific requirements to germinate. In contrast, seeds exhumed from the flooded lowland germinated ca. 65% when nitrates were combined with red light pulses, significantly higher than in the rest of the treatments. Seeds exhumed from drained mesocosms germinated equally in all treatments. However, in the seeds exhumed from the flooded mesocosms, nitrates increased germination by more than 20% compared with seeds imbibed in water. Seeds germinated ca. 85% when nitrates were combined with red light pulses, significantly higher than in the other treatments. We can conclude that after flooding, S. parviflora seeds require nitrate and light to germinate. Therefore, a large fraction of seeds do not germinate unless nitrates are combined with light, indicating a drawdown period after floods and vegetation gaps.     

Germination requirements inCarex kobomugi (Sea Isle)

Laboratory and field germination experiments inCarex kobomugi seeds were pursued to clarify their germination requirements and availability of the requirements in the field. In the laboratory experiments, more than 50% of the seeds ofC. kobomugi germinated under 35/30C or 25/20C when they were scarified with 98% H₂SO₄ after removal of their utricles, and chilled in moist condition for 28 to 42 d. Seeds with utricles or those without scarified with H₂SO₄ did not germinate. Seeds sown at 10-cm depth at the Kado-ori coast on 11 February 1991 after soaked in H₂SO₄ showed 40% germination by 29 April 1991, whereas those without H₂SO₄ treatment did not germinate. These results suggest that seed-coat impermeability and embryo immaturity are possible causes of the dormant state in seeds ofC. kobomugi ripen in summer. In the field, the moist-chilling condition is available in winter and the seeds can germinate in the following spring if the seed-coat impermeability is relaxed before winter.     

Morphophysiological dormancy in seeds of two North American and one Eurasian species of Sambucus (Caprifoliaceae) with underdeveloped spatulate embryos

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.     

The germination characteristics of <Emphasis Type="Italic">Scrophularia marilandica</Emphasis> L. (Scrophulariaceae) seeds

Investigations on seeds of Scrophularia marilandica L. were undertaken to determine their germination requirements. Seeds were collected from three naturally occurring sites and one greenhouse-grown population in London, Ontario in September and October of 1997. Some were set to germinate immediately after collection; others were stored in or on soil outside and/or under controlled laboratory conditions before testing. Germination was assessed under two light/temperature regimes (35°C 14 h light, 20°C 10 h dark and 25°C 14 h light, 10°C 10 h dark), in continuous darkness, and in the presence of two germination-promoting chemicals (GA₃ and KNO₃). Fresh seeds germinated best at 35/20°C, while stored seeds germinated best at 25/10°C. No differences in percent germination were found among three seed-maturity stages. All chemical treatments, except 0.01 M KNO_3, increased percent germination. Significant differences were found both among and within sites for most chemical treatments, but exposure to 3 × 10⁻⁴ M GA₃ caused almost every seed to germinate. When compared to the control, both the gibberellic acid and the soil-storage treatments contributed to faster germination. Exposure of seeds to naturally prevailing conditions on the soil surface followed by testing under the 25/10°C regime produced the highest percent germination. No seeds germinated in the dark. In summary, seeds of S. marilandica exhibit physiological dormancy, which can be alleviated by exposure to light, after-ripening and/or cold stratification. It is probable that the differences in germination response among sites can be attributed to differences in environmental conditions during seed production. These experiments indicate that the seeds of S. marilandica must be buried shortly after dispersal in order to form a persistent seed bank.     

Germination responses of Discopodium penninervium Hochst. to temperature and light

The germination responses of Discopodium penninervium were tested at different constant and alternating temperature regimes as well as under various light conditions both in the laboratory and glasshouse. Seeds incubated at 10, 15, 20, 25 and 30 °C failed to germinate. When the seeds were incubated at alternating temperatures of 20/12 °C and 30/12 °C under continuous light, germination was 89 and 61%, indicating that the species requires alternating temperatures as a cue for germination. However, germination declined as the amplitude of alternating temperatures increased from 8 °C and was completely inhibited at an amplitude of 23 °C, suggesting that the optimum amplitude is around 8 °C. Germination was less than 10% in light and nil in darkness at 20 °C in the laboratory. In contrast, seeds incubated at 20/12 °C germinated to 96 and 86% in light and darkness, respectively. Seeds incubated under leaf shade in the glasshouse failed to germinate whereas those incubated under direct daylight and darkness germinated to 44 and 50%, respectively, 30 days after sowing. When seeds incubated under leaf shade and in darkness were exposed afterwards to light, final percent germination was 83% from seeds incubated initially under direct daylight, 79% from those incubated under leaf shade and 86% from those incubated in darkness. The requirement for alternating temperatures and light rich in red:far red ratio to break the dormancy of seeds of D. penninervium could restrict germination to gaps in the vegetation. The results conform with the ecology of the species.     

DORMANCY IN SEEDS OF FRASERA CAROLINIENSIS (GENTIAN ACEAE)

In freshly matured seeds of the long-lived monocarpic perennial, Frasera caroliniensis Walt., the embryos are underdeveloped and physiologically dormant. Dormancy was broken by a long period of stratification (chilling) at 5 C. Seventy six percent of the seeds germinated at 20 C (day)/10 C (night) after 98 days of chilling at 5 C, while seeds kept at 5 C germinated to 87% after 205 days. A warm, moist pretreatment was not required for subsequent breaking of dormancy at 5 C. Embryos in fresh seeds averaged 1.3 mm long, but after 12 weeks of chilling they averaged 4.1 mm. Thus, the embryos require a period of chilling to become fully developed, after which seeds can germinate at the afterripening temperatures (5 C) or at some higher temperature. Seeds of F. caroliniensis fit Nikolaeva's (1977) morpho-physiological complex dormancy type.     

GERMINATION ECOPHYSIOLOGY OF ARENARIA GLABRA,A WINTER ANNUAL OF SANDSTONE AND GRANITE OUTCROPS OF SOUTHEASTERN UNITED STATES

The germination ecophysiology of Arenaria glabra Michx., a characteristic winter annual plant species of granite and sandstone outcrops of southeastern United States, was investigated. Seeds germinate in early autumn, plants overwinter in the rosette stage and then flower, set seeds, and die in late spring; seeds are dispersed soon after maturity. Eighty-five to 90% of freshly-matured seeds were innately dormant, and the other 10–15% germinated only at temperatures lower than those that occur in the habitat at the time of seed dispersal in June. During the summer after-ripening period, seeds stored dry under ambient laboratory conditions exhibited progressive increases in rates and total percentages of germination, a widening of the temperature range for germination, and a loss of the light requirement. At a 14-hr daily photoperiod, seeds kept on continuously moist soil germinated to 83% at simulated July and August temperatures during July and August, and the remainder germinated at September temperatures in September. On the other hand, seeds subjected to alternate wetting and drying during July and August germinated to only 9% during those 2 months, and the remainder germinated after the soil was kept continuously moist, beginning on 1 September, at simulated habitat temperatures during September and October. Thus, the timing of germination of A. glabra in the field is controlled by an interplay of the seeds' physiological state with the dynamics of temperature and soil moisture conditions.     

Palms Use a Bluffing Strategy to Avoid Seed Predation by Rats in Brazil

The goal of this study was to ascertain why the production of variable seediness is advantageous for Attalea phalerata palms. Our hypothesis was that variation reduces seed predation by the spiny rats Thrichomys pachyurus and Clyomys laticeps. Although there is a positive correlation between endocarp size and number of seeds, endocarps sometimes contain more or fewer seeds than expected; palms bluff about the number of seed per endocarp. Therefore, rats do not know how many seeds an endocarp contains. To model rats' predating behavior, we applied Charnov's Marginal Value Theorem. The model shows that rats attack endocarps only when the energy gain is higher than the energy available in the habitat. Hence, it is not advantageous to eat all the seeds inside an endocarp. This explains why 45 percent of forest endocarps and 35 percent of savanna endocarps were still viable after predation. We then applied the model to two simulated endocarp populations with less variability in the number of seeds per endocarp size and determined that viable diaspores after predation were reduced to 15 percent. With less variability, palms cannot bluff about the number of seeds inside endocarps and predators can predict accurately how many seeds they should try to eat. Uncertainty about the number of seeds diminished predation but gave selective advantage to multiseeded fruits. Therefore, the bluffing strategy would be evolutionarily stable only if it were counterbalanced by other forces. Otherwise, predators would win the bluffing game. Abstract in Portuguese is available at http://www.blackwell-synergy.com/loi/btp .     

Sensitivity of seed germination to water stress in high-altitude populations of a threatened palm species

• Divergence in seed germination patterns among populations of the same species is important for understanding plant responses to environmental gradients and potential plant sensitivity to climate change. In order to test responses to flooding and decreasing water potentials, over 3 years we germinated and grew seeds from three habitats of Euterpe edulis Mart. occurring along an altitudinal gradient.
• Seed germination and root growth were evaluated under different water availability treatments: control, flood, −0.4 MPa, −0.8 MPa, in the years 2012, 2013 and 2014, and in the final year of the experiment (2014) at −1.0 MPa and −1.5 MPa.
• Seeds from the montane habitat did not germinate in the flooding treatment. Seed germination of all three habitats decreased in the −1.5 MPa treatment and the montane habitat had lowest germination in this treatment. Time required for half of the seeds to germinate increased up to −0.8 MPa. Seeds from montane habitats germinated more slowly in all treatments. The only difference in seed germination synchrony was an increase in the submontane population under the flooding treatment. However, synchrony decreased at the lowest water potentials. Roots of the montane population were more vigorous in most treatments, except at −0.8 MPa.
• The unusual ability of these seeds to germinate at low water potentials might be related to early seed germination at the onset of the rainy season, which potentially decreases seed predation pressure. Seeds of the montane population were more sensitive to both types of water stress. A predicted increase in the frequency and intensity of extreme high rainfall or drought events may predispose early stages of this population to adverse factors that might negatively affect population viability with elevational in future climate change scenarios.