Dormancy cycles in buried seeds of three perennial Xyris (Xyridaceae) species from the Brazilian campo rupestre

• Dormancy cycles are an important mechanism for avoiding seed germination under unfavourable periods for seedling establishment. This mechanism has been scarcely studied in tropical species. Here, we studied three tropical and perennial species of Xyris, X. asperula, X. subsetigera and X. trachyphylla, to investigate in situ longevity and the existence of seasonal seed dormancy cycles.
• Seeds of three species of Xyris were buried in their natural habitat, with samples exhumed bimonthly for 18 months. Germination of exhumed seeds was assessed under a 12‐h photoperiod over a broad range of temperatures. Seeds of X. trachyphylla were also subjected to treatments to overcome secondary dormancy.
• Seeds of all species are able to form a persistent seed bank and exhibit seasonal changes in germinability. Secondary dormancy was acquired during the rainy summer and was overcome during the subsequent dry season (autumn/winter). Desiccation partially overcomes secondary dormancy in X. trachyphylla seeds.
• Soil seed bank persistence and synchronisation of seed germination under favourable conditions for seedling establishment contribute to the persistence and regeneration of X. asperula, X. subsetigera and X. trachyphylla in their natural environment.

     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

Dormancy cycling and persistence of seeds in soil of a cold desert halophyte shrub

Background and Aims

Formation of seed banks and dormancy cycling are well known in annual species, but not in woody species. In this study it was hypothesized that the long-lived halophytic cold desert shrub Kalidium gracile has a seed bank and dormancy cycling, which help restrict germination to a favourable time for seedling survival.

Methods

Fresh seeds were buried in November 2009 and exhumed and tested for germination monthly from May 2010 to December 2011 over a range of temperatures and salinities. Germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were investigated in the field.

Key Results

Seeds of K. gracile had a soil seed bank of 7030 seeds m⁻² at the beginning of the growing season. About 72 % of the seeds were depleted from the soil seed bank during a growing season, and only 1·4 % of them gave rise to seedlings that germinated early enough to reach a stage of growth at which they could survive to overwinter. About 28 % of the seeds became part of a persistent soil seed bank. Buried seeds exhibited an annual non-dormancy/conditional dormancy (ND/CD) cycle, and germination varied in sensitivity to salinity during the cycle. Dormancy cycling is coordinated with seasonal environmental conditions in such a way that the seeds germinate in summer, when there is sufficient precipitation for seedling establishment.

Conclusions

Kalidium gracile has three life history traits that help ensure persistence at a site: a polycarpic perennial life cycle, a persistent seed bank and dormancy cycling. The annual ND/CD cycle in seeds of K. gracile contributes to seedling establishment of this species in the unpredictable desert environment and to maintenance of a persistent soil seed bank. This is the first report of a seed dormancy cycle in a cold desert shrub.     

Branch direction in Carnegiea gigantea (Cactaceae): Regional patterns and the effect of nurse plants

Abstract. Branch direction in Carnegiea gigantea (saguaro) cacti affects PAR interception and CO₂ uptake, and may influence reception of photomorphogenic cues. Branch direction data were collected in 29 saguaro populations over Arizona to determine if the southerly bias observed in previous studies at two locales can be generalized to the American range of the saguaro, and to determine the effect of shading by nurse plants on branch direction. Locally observed S bias from previous studies can be extended and generalized to include the American saguaro range. Branches were statistically uniform across environmental variability regarding temperature and precipitation. The S bias was not found in saguaros shaded by large nurse plants, supporting the assertion that branch direction occurs, in part, to maximize receipt of insolation.     

果翅对梭梭属(Haloxylon)种子萌发行为的调控

种子萌发行为的受控机制是生殖生态学的重要研究内容。通过果翅存留实验研究了在不同贮藏时期果翅对梭梭和白梭梭种子萌发的影响。结果表明：①新成熟的梭梭种子具有高的萌发率（〉90％）。梭梭果翅对秋天新成熟的种子萌发有显著的抑制作用（萌发率〈50％）,使种子处于强迫休眠状态;果翅对种子萌发的抑制作用随着贮藏时间的推移逐渐降低,到翌年春天（4月）这种抑制作用已完全解除,果翅对梭梭种子的抑制主要是化学抑制。果翅对梭梭种子萌发行为的调控作用确保了种子在合适的时间萌发与种群的成功定居,也正是梭梭在荒漠地区广泛分布的主要原因。②新成熟的自梭梭种子萌发率为59．5％,贮藏1个月后达到80％。表明种子存在短时期的生理后熟现象。白梭梭果翅在每一贮藏时期对其种子的萌发都具有显著的抑制作用,翅＋种子（将果翅与种子剥离。一起放人培养皿中）也部分地抑制种子的萌发,因此,果翅对白梭梭种子萌发的抑制既有化学抑制也存在机械抑制。果翅对种子萌发的抑制,限制了白梭梭的分布。     

Decomposition rate of ocotillo (Fouquieria splendens) wood in the desert of southern California and its use in estimating adult survival by life-cycle graph analysis

Dead and downed ocotillos (Fouquieria splendens) are common in the Colorado Desert of southern California and pose a question concerning the life-span of adult plants that have been reported to be long-lived: if ocotillo is long-lived, decay rates must be very slow. Our study focused on decomposition and disintegration of above-ground ocotillo wood over 14 years to obtain a rate of change in dead mass. Disintegration rate was used in a new application of life-cycle graph analysis to calculate persistence time of carcasses, which in turn was used to estimate adult survival rates. We conclude that previous estimates of life spans may be too conservative and half-life of adult shrubs is over 200 years. Our approach to estimating adult survival from decay rates provides an independent test of survival transitions based on live plants for some species and so is an additional tool for developing life cycle models.     

榛树种子的休眠和萌发

鉴定榛子各部分浸提液的结果显示 ,果皮、种皮及胚都存在导致休眠的抑制物质 ;榛树果实有一定的吸水性 ,据此推测种子休眠可能不是因果皮的不透水性造成 ;0 .5、2、5mg·L-16 BA可以解除种子休眠 ,促进萌发。     

Comparative longevity of Australian orchid (Orchidaceae) seeds under experimental and low temperature storage conditions

Seeds from ten terrestrial orchid species, nine from the south‐west Australian biodiversity hotspot (Caladenia arenicola, Caladenia flava, Caladenia huegelii, Diuris laxiflora, Microtis media ssp. media, Pterostylis recurva, Pterostylis sanguinea, Thelymitra crinita and Thelymitra macrophylla) and one from south‐east Australia (Diuris fragrantissima), were placed into experimental storage to assess their relative longevity and likely optimal conditions for long‐term conservation seed banking. Seeds from all species were desiccation tolerant, germinating after drying at 23% relative humidity (C. arenicola, C. huegelii, P. sanguinea and T. macrophylla) or 5% relative humidity (C. flava, D. laxiflora, M. media ssp. media, P. recurva and T. crinita) at 23 °C. From automatedly determined moisture adsorption and desorption isotherms at 23 °C, these equate to tolerance of drying to 0.03–0.06 g water g^?1 dry weight or 0.013–0.028 g water g^?1 dry weight, respectively. Results of storage experiments at a range of moisture contents and temperatures suggest conventional seed bank storage at ?18 °C after equilibration at c. 23% relative humidity (at 23 °C) may be suitable for most of the species, although there was higher germination of P. recurva seeds stored at ?80 °C and of M. media ssp. media seeds equilibrated at 75% relative humidity. However, there was considerable variation in germination of seeds sampled after different storage periods, making it difficult to identify optimal storage conditions definitively. Results of comparative longevity storage experiments at 60% relative humidity and 40 °C suggest seeds from these orchid species are short‐lived compared with non‐orchid species, and with Australian species in particular. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 26–41.     

脱落酸调控种子休眠和萌发的分子机制

脱落酸(ABA)是调控种子休眠和萌发过程的主要植物激素。种子内源ABA含量和种胚对ABA敏感性共同调控种子休眠和萌发过程, 确保植物种子以休眠状态在逆境中保持其自身繁衍能力, 并在适宜的环境下启动萌发程序。种子ABA合成代谢和ABA信号转导途径涉及许多重要基因家族, 它们通过复杂的调控网络精确地控制着种胚发生、种子成熟、休眠及萌发进程。该文对ABA调控种子休眠和萌发的分子机制最新研究进展进行综述, 并展望了今后的研究方向。     

盐生草(Halogeton glomeratus)二型种子的休眠与萌发

盐生草的果实和种子存在二型性,这两种类型的种子在形状、大小、颜色及包被其花被片背部是否具翅上均有显著差异.绿色种子,圆形,直径为(1.552±0.116) mm,宿存花被革质,背部有紫红色翅状附属物,单粒重为(0.808±0.033)mg;黄色种子,椭圆形,长为(1.752±0.155) mm,宽为(1.146±0.088) mm,宿存花被革质,背部无翅状附属物,单粒重为(0.568±0.011) mg.两种种子在3个变温条件(5/25℃、5/25℃、15/25℃,暗12 h /光12 h)下的萌发率均较低,绿色种子为36%,而黄色种子为17%(15℃/25℃).延长储藏时间和划破种皮均能显著提高绿色种子的萌发率,表明绿色种子属于非深度生理休眠.随着储藏时间的延长,黄色种子的萌发率也能缓慢提高,但不显著,而划破种皮能够显著促进其萌发,表明黄色种子属于深度生理休眠.     

Demographic patterns of Ferocactus cylindraceus in relation to substrate age and grazing history

Three subpopulations of Ferocactus cylindraceus, a short-columnar cactus of the Sonoran and Mojave deserts, were sampled in Grand Canyon, Arizona, USA, at sites representing a range of substrate ages and different grazing histories. Age-height relations were determined from annual growth, then used to estimate probable year of establishment for each cohort. Eight years between 1944 and 1992 were especially favorable for establishment. Six of these 8 years coincided with El Niño-Southern Oscillation conditions, indicating that as for many woody plants in arid regions, somewhat unusual climatic conditions are necessary if populations are to replace themselves.Comparison of age structures showed that established and developing populations have somewhat different dynamics in that the rate of population increase was slowest on the youngest terrace. On the ancient terraces, about half the plants were less than 25 years old. Plants older than 40 years were few; however the oldest plants in the study (about 49 years) grew on the ancient terraces. On the recent terrace, 76% of the subpopulation was 25 years or younger, and the oldest living plant was about 36 years of age. The age structures of subpopulations on grazed and ungrazed sites also differed markedly. On ungrazed sites, subpopulations were more or less at equilibrium, with enough young plants to replace old ones as they died. In contrast, the subpopulation on the grazed site was in a state of marked disequilibrium. Grazing before 1981 largely extirpated a palatable subshrub that was probably an important nurse plant. Until the shrub population at Indian Canyon recovers from decades of burro grazing, a rebound in F. cylindraceus establishment is not to be expected.Nomenclature: Hickman, J. C. (ed.) 1993. The Jepson manual: Higher plants of California. University of California Press, Berkeley.     

Digestion time and intactness of seeds ingested by Sturnira lilium (E. Geoffroy, 1810) (Mammalia,Chiroptera)

Bats are responsible for many ecological services, such as seed dispersal of several plant species, contributing to the processes of succession and forest regeneration. A factor that can interfere with this process is the animal digestion, which can affect germination, altering the patterns of seedling distribution. The effects of seed passage through bats’ guts varies with the species, leading to some discrepancies in the literature. In this study, we tested the digestion time of one Phyllostomidae bat species, Sturnira lilium, in two Neotropical plants: Solanum paniculatum and Ficus organensis, and the effects on seed germination. The experiment was conducted in captivity and the germination tests were made in laboratory conditions. The results suggested that most seeds ingested by S. lilium are dispersed within 40 min for both species and the digestion seems not to significantly affect the germination of F. organensis, despite the slight acceleration of germination. In S. paniculatum, germination occurred only in the control (39%), whereas in the treatments, all the seeds remained dormant during the 25 experimental days. In this case, the digestion of S. lilium possibly contributes to the formation of seed banks, randomizing the temporal distribution of seedlings.     

野生鸭儿芹种子休眠特性及破除方法

鸭儿芹种子具有休眠特性,且休眠期长,不经任何处理的种子很难萌发,影响其人工种植。研究了鸭儿芹种子的休眠特性和解除休眠的最佳方法,为我国人工种植野生鸭儿芹提供理论依据。结果表明:TTC法对种子活力的测定表明有活力的种子为(55.33±3.71)%;切破种皮种子与完整种子吸水率在前12 h相差较大,但最终吸水率相差不大,分别达到(70.00±1)%和(68.32±0.32)%,表明种皮并不阻碍种子吸水;种子中存在内源抑制物,其粗提液在较低浓度下即可抑制芹菜种子的萌发;鸭儿芹种子成熟时胚未分化完全,胚率为(28.65±2.488)%,经过低温处理后完成后熟,胚率达到(65.93±3.86)%,萌发率达到100%,因此鸭儿芹种子具有形态生理休眠特性。清水浸种和低温冷藏共同处理可有效解除其休眠,浸种和低温冷藏具有交互效应,浸种36 h、5℃冷藏30d即可解除其休眠,萌发率达到100%,发芽势达到(91.11±0.91)%。已破除休眠的种子适宜其萌发的温度范围扩大(15.0—27.5℃),而且在土壤中也可较好地萌发,萌发率达到(96.67±3.33)%,发芽势达到(71.11±1.93)%。     

Field observations of intercolony aggression and territory changes inHeterotermes aureus (Isoptera: Rhinotermitidae)

Intercolony aggression was observed for a southwestern desert subterranean termite, Heterotermes aureus(Snyder), in bait plots in a Sonoran Desert grassland in Arizona. Intermittent conflict took place at the sites of corrugatedpaper rolls on the boundary between two colonies. Rows of dead termites locked in combat were found within the corrugations, with additional dead termites usually in the soil beneath the roll. Dead soldiers constituted 64 to 100% of termites recovered from sites of intercolony conflict. Ongoing intercolony aggression is likely involved in the maintenance of discrete H. aureusterritories.     

The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.     

Burial of canopy-stored seeds in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) and its ecological significance

Agriophyllum squarrosum Moq. is a dominant annual on sand dunes in the arid regions of central Asia. A high percentage of seeds is retained on dead plants which become covered by moving sand, but little is known about the ecological significance of burial of canopy-stored seeds. We investigated the size and dynamics of the buried canopy-stored seed bank and effects of burial on seed germination. In March (during the windy season), May (beginning of the germination season), and July (middle of the growing season), the number of seeds per square meter in sample plots in the dunes was 623, 223 and 22, respectively, with 54.6, 30.6 and 12.9% of the total seeds retained on buried plant canopies. In a controlled experiment, more seedlings emerged from released (dispersed) than from canopy-stored seeds when burial depth was the same. No viable ungerminated released seeds were found, but 45–80% of the ungerminated canopy-stored seeds were viable. In general, with an increase in applied water germination of released seeds buried at a depth of 1 or 2 cm and of canopy-stored seeds buried at 1 cm increased, but regardless of watering regime few or no released seeds at 4 cm or canopy-stored seeds at 2 or 4 cm germinated. Significantly more seedlings emerged from plants buried in a horizontal than in a vertical position. Seedlings originating from buried canopy-stored seeds on an active dune accounted for only 5.4% of the total seedlings emerging, and most of them emerged later than those from released seeds. Thus, seed release is more effectively postponed in buried than in exposed canopies, and burial of canopy-stored seeds is a mechanism that helps regulate seed germination and seedling emergence of A. squarrosum on active dunes.     

Seed hydration memory in Sonoran Desert cacti and its ecological implication

Cactus seeds on the soil surface in the desert are subjected to periods of drought that last for up to a few months, and thus they are typically under discontinuous hydration (or discontinuous dehydration). Apparently, they can tolerate long periods of dehydration after single or multiple hydration events and subsequently germinate in accordance with the previous hydration experience. This was verified in three cactus species from the Sonoran Desert. Seeds of Stenocereus thurberi hydrated for 72 or 80 h followed by a dehydration period lasting for 4, 14, 70, 120 or 181 d germinated 2–3 d earlier and had 1.4–2 times shorter mean germination time (MGT) than untreated seeds. Seeds given shorter hydration periods also began to germinate sooner than the controls. MGT was shorter only when the hydration period was 48 h or longer. Final germination percentages were not affected by these treatments, only the MGT. Except for differences in germination percentages, similar results were found for Pachycereus pecten-aboriginum and Ferocactus peninsulae. When the cycle of 24 h hydration followed by 4 d dehydration was repeated one or two times, the effect was cumulative: MGT was equal to 48 and 72 h hydration, respectively. These results suggest a phenomenon of “seed hydration memory,” the ability of seeds to retain during dehydration periods those physiological changes that result from seed hydration. Thus, treated seeds subsequently germinated earlier then untreated seeds, regardless of the duration of dehydration period. This led to a greater biomass accumulation and thus to higher survival in seedlings from treated than from untreated seeds.     

Effects of brassinosteroids on conditioning and germination of clover broomrape (Orobanche minor) seeds

Brassinolide [2, 3, 22R,23R-tetrahydroxy-24S-methyl-B-homo-7-oxa-5--cholestan-6-one] and its related compounds, brassinosteroids, applied at the early stages of conditioning shortened the conditioning period required before clover broomrape seeds would germinate after exposure to germination stimulants, such as dl-strigol and natural stimulants from red clover (Trifolium pratense L.) root exudate. Brassinosteroids applied after conditioning increased the rate of the seed germination induced by these stimulants. The inhibitory effect of light on seed germination when it was induced by dl-strigol could be overcome by brassinosteroids. Brassinosteroids also eliminated the inhibitory effects of light and dl-strigol applied at the early stages of conditioning. GA₃ was also effective in causing seed conditioning and increased the rate of the germination induced by these stimulants. There was a relationship between brassinosteroids and GA₃ in many of the experiments conducted. These findings may have practical implications in increasing the effectiveness of applying germination stimulants in the field to soils for suicidal germination of broomrape seeds.     

Testing myrmecochory from the ant’s perspective: The effects of Datura wrightii and D. discolor on queen survival and brood production in Pogonomyrmex californicus

Interspecific interactions are often assumed to be mutualistic if one species appears to benefit. However, most studies do not test whether both participants benefit. Myrmecochory, or seed dispersal by ants, is characterized by a lipid-rich appendage, or elaiosome, on a seed. Typically, ants gather the diaspores (i.e., seeds with elaiosomes), carry them to the nest, consume the elaiosome, and discard the seed unharmed either inside the nest or on a refuse pile. The benefit to the ants is presumably the nutritional content of the elaiosome, whereas benefits to the plant include dispersal from the parent plant, protection from predators, reduced seedling competition, protection from fire, or transportation to nutrient-rich microsites. Most studies of myrmecochory focus on potential benefits to the plants and simply assume that ants receive a benefit from consuming elaiosomes. I tested whether Pogonomyrmex californicus benefits from consuming Datura wrightii and D. discolor elaiosomes by raising newly-mated queens (i.e., foundresses) on different diets and measuring their survival and brood production. Foundresses reared solely on D. wrightii or D. discolor had similar probabilities of surviving and producing brood as foundresses fed a standard diet, but the number and developmental stage of the brood produced was severely reduced. Because the initial number of brood produced is critical for successful colony establishment, the future fitness of foundresses consuming only Datura is likely reduced. In addition, adding Datura to a standard diet did not increase queen survival or brood production. Although it is possible that Datura may help sustain a colony through periods of scarcity, P. californicus do not appear to receive nutritional benefits from myrmecochorous interactions with Datura in the northern Sonoran Desert. Received 4 July 2005; revised 17 April 2006; accepted 9 May 2006.