Ecological genetics of seed germination regulation in Bromus tectorum L.

Regulation of seed germination phenology is an important aspect of the life history strategy of invading annual plant species. In the obligately selfing winter annual grass Bromus tectorum, seeds are at least conditionally dormant at dispersal in early summer and lose dormancy through dry-afterripening. Patterns of germination response at dispersal vary among populations and sometimes across years within populations. To assess the relative contribution of genotype and maturation environment to this variation, we grew progeny of ten parental lines from each of six contrasting populations in a common greenhouse environment. We then tested the germination responses of recently harvested seeds of the putative full-sib progeny at five incubation temperatures. Significant germination response differences among populations were observed in greenhouse cultivation, and major differences among full-sib families were evident for some populations and traits. Among-population variation accounted for over 90% of the variance in each trait, while within-family variance accounted for 1% or less. Germination responses of greenhouse-grown progeny were positively correlated with the responses of wild-collected seeds, but there was a tendency for lowered dormancy at higher incubation temperatures. This tendency was more marked in populations from cold desert, foothill, and plains habitats, suggesting a genotype-maturation environment interaction. Differences among populations in the amount of among-family variance were more evident at lower incubation temperatures, while among-family variance was more uniformly low at summer incubation temperatures. Populations from predictable extreme environments (subalpine meadow and warm desert margin) showed significantly less variation among families than populations from less predictable cold desert, foothill, and plains environments. Low among-family variance was not specifically associated with small population size or marginality of habitat, as small marginal populations from unpredictable environments showed variance as high as that of large populations. In populations with high among-family variance for germination traits, germination responses tended to be correlated across incubation temperatures, making it possible to characterize families in terms of their general dormancy status. The results indicate that seed germination regulation in this species is probably under strong genetic control, and that habitats with temporally varying selection are occupied by populations that tend to be more polymorphic in terms of their germination response patterns. Received: 19 May 1998 / Accepted: 27 January 1999     

Ecological genetics of Bromus tectorum

Summary By incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.     

Ecological genetics of vernalization response in Bromus tectorum L. (Poaceae)

BACKGROUND AND AIMS: Bromus tectorum (cheatgrass or downy brome) is an exotic annual grass that is dominant over large areas of former shrubland in western North America. To flower in time for seed production in early summer, B. tectorum plants generally require vernalization at winter temperatures, either as imbibed seeds or as established seedlings. METHODS: Variation in response to increasing periods of vernalization as seeds or seedlings for progeny of ten full-sib families from each of four B. tectorum populations from contrasting habitats was studied. KEY RESULTS: As vernalization was increased from 0 to 10 weeks, the proportion of plants flowering within 20 weeks increased, weeks to initiation of flowering decreased, and seed yield per plant increased, regardless of whether plants were vernalized as seeds or seedlings. Most of the variation was accounted for by differences among populations. Plants of the warm desert population flowered promptly even without vernalization, while those of the cold desert, foothill and montane populations showed incremental changes in response variables as a function of vernalization period. Populations differed in among-family variance, with the warm desert population generally showing the least variance and the cold desert population the most. Variation among populations and among families within populations decreased as vernalization period increased, whereas the non-genetic component of variance showed no such pattern. CONCLUSIONS: Variation in vernalization response was found to be adaptively significant and apparently represents the result of contrasting selection regimes on a range of founder genotypes.     

Patterns of seed after-ripening in Bromus tectorum L

For grass seeds that lose dormancy through after ripening indry storage, the probability of germination following a particularwetting event can be predicted only if the relationship betweenstorage temperature and change in after-ripening status is known.This study examined patterns of seed dormancy loss in Bromustectorum L., quantifying changes in germination percentage,speed, and uniformity through time. Seed collections from threesemi-arid habitats were stored at temperatures from 10–40C. At monthly intervals, subsamples were incubated at 5/15,10/20, 15/25, and 20/30 C. For recently harvested seeds, germinationpercentage, mean germination time, and days between 10% and90% of total germination (D90–D10) ranged from 1–75%,10–24 d, and 10–20 d, respectively. Recently harvestedseeds were generally most dormant, slowest to germinate andleast uniform at high incubation temperatures. In contrast,after ripened seeds for all collections had nearly 100% germination,mean germination times <5 d, and D90–D10 values <5d. Three indices were used to characterize after-ripening ratesfor each seedlot at each incubation temperature. The mean dormancyperiod, the mean rate index, and the mean uniformity index definedthe storage period required for seedlots to become half as dormantas at harvest, to progress half-way to the fastest speed, andto progress half-way to the greatest uniformity, respectively.Seeds required longer storage to germinate uniformly than togerminate completely or quickly, because germination time-coursecurves for incompletely after-ripened seeds were positivelyskewed rather than sigmoidal. Mathematically, the three indiceswere described as negative exponential functions of storagetemperature, which suggests that after-ripening is likely completedin late summer or early autumn regardless of summer conditions. Key words: Seed dormancy, germination timing     

Fire promotes downy brome (Bromus tectorum L.) seed dispersal

Particularly well-known among the many impacts of the invasive annual grass downy brome (Bromus tectorum, Poaceae) is its ability to alter fire cycles and increase in abundance after fire. However, little is known about how fire influences B. tectorum dispersal. We quantified fire effects on B. tectorum dispersal using three recently burned areas in the western region of the Colorado Rocky Mountains by marking diaspores (seeds) with fluorescent powder, and then recovering them at night using ultraviolet lights. Diaspores were of two types: with and without sterile florets attached. We also characterized vegetation cover and near-surface wind speed in burned and unburned areas. Diaspores travelled much farther in burned areas than in nearby unburned areas (mean ± standard error at the end of the experiment: 209 ± 16 cm and 38 ± 1 cm, respectively; maximal distance at the end of the experiment: 2,274 cm and 150 cm, respectively), indicating an increase in dispersal distance after fire. Diaspores with sterile florets attached dispersed longer distances than those without sterile florets (mean ± standard error at the end of the experiment: 141 ± 14 cm and 88 ± 7 cm, respectively). Vegetation cover was lower and wind speeds were higher in the burned areas. Our results indicate that at least one of the mechanisms by which the spread of B. tectorum is promoted by fire is through increased seed dispersal distance. Preventing movement of seeds from nearby infestations into burned areas may help avoid the rapid population expansion often observed.     

旱雀麦(Bromus tectorum L.)入侵机制的研究进展

旱雀麦(Bromus tectorum)在世界上广泛分布,但在北美却是一种入侵能力较强的物种,对当地的植物群落结构组成和生态系统功能产生重要影响.本文从旱雀麦的生境和扩散路径、本身的生物学特性、其与新栖息地土著物种之间的相互作用、新栖息地的环境变化对旱雀麦的影响等方面综述了近年来关于旱雀麦入侵机制的研究进展.研究表明,影响旱雀麦入侵的因素是多样的,阐明旱雀麦的入侵机制对于推动生态学的理论发展和控制旱雀麦的大规模入侵具有重要意义.     

A simulation model to predict seed dormancy loss in the field for Bromus tectorum L

Bromus tectorum L. (cheatgrass) is an invasive winter annual whose seeds lose dormancy through dry after-ripening. In this paper a thermal after-ripening time model for simulating seed dormancy loss of B. tectorum in the field is presented. The model employs the hydrothermal time parameter mean base water potential (b(50)) as an index of dormancy status. Other parameters of the hydrothermal time equation (the hydrothermal time constant HT, the standard deviation of base water potentials b, and the base temperature Tb) are held constant, while b(50) is allowed to vary and accounts for changes in germination time-course curves due to stage of after-ripening or incubation temperature. To obtain hydrothermal time parameters for each of four collections, seeds were stored dry at 20C for different intervals, then incubated in water (O MPA) or polyethylene glycol (PEG) solutions (-0.5, -1.0, -1.5 MPa) at 15 and 25C. Germination data for the thermal after-ripening time model were obtained from seeds stored dry in the laboratory at 10, 15, 20, 30, 40, and 50°C for 0 to 42 weeks, then incubated at two alternating temperatures in water. Change in b(50) was characterized for each collection and incubation temperature as a linear function of thermal time in storage. Measurements of seed zone temperature at a field site were combined with equations describing changes in b(50) during after-ripening to make predictions of seed dormancy loss in the field. Model predictions were compared with values derived from incubation of seeds retrieved weekly from the field site. Predictions of changes in b(50) were generally close to observed values, suggesting the model is useful for simulating seed dormancy loss during after-ripening in the field.     

Seed aging,delayed germination and reduced competitive ability in Bromus tectorum

In annual plants, increased competitive advantage has often been attributed to rapid germination and early establishment. In contrast, many annual species exhibit some degree of delayed germination (i.e., seed dormancy) that results in the formation of age structure within the seed population. Delayed germination can be an effective bet-hedging strategy in variable or unpredictable environments as a seed bank can buffer against years with reproductive failures and reduce the probability of local extinction. However, there has been little consideration of the direct effects of aging within the seed pool although the potential demographic costs of such a strategy (e.g., mortality in the seed bank or delayed reproduction) are well known. We used aged (4 year-old) and freshly produced seed from meadow steppe and sagebrush steppe populations of an annual grass (Bromus tectorum)to investigate the importance of seed age on seedling vigor and competitive ability. Aged seed from the meadow steppe population exhibited delays in germination that reduced plant growth and final biomass when the plants were grown with competition. Aged seed from the sagebrush steppe population did not exhibit delays in germination. By including a treatment that experimentally delayed the germination of freshly produced meadow steppe seed, we also examined the effects of delayed germination alone. A comparison of results from this delay treatment with those from the aged seed treatment suggested that the reduced competitive ability of meadow steppe plants produced from aged seed, although largely a result of the temporal delay in germination, was partly due to reduced seed vigor. Together these results indicate that physiological costs associated with seed age may affect aboveground competitive interactions and, in turn, the relative fitness of older cohorts in the soil seed bank.     

Introduction history and population genetics of the invasive grass Bromus tectorum (Poaceae) in Canada

The invasive annual Bromus tectorum (cheatgrass) is distributed in Canada primarily south of 52° N latitude in two diffuse ranges separated by the extensive coniferous forest in western Ontario. The grass was likely introduced independently to eastern and western Canada post-1880. We detected regional variation in the grass's genetic diversity using starch gel electrophoresis to analyze genetic diversity at 25 allozyme loci in 60 populations collected across Canada. The Pgm-1a & Pgm-2a multilocus genotype, which occurs in the grass's native range in Eastern Europe, is prevalent in eastern Canada but occurs at low frequency in western Canada. In contrast, the Got-4c multilocus genotype, found in the native range in Central Europe, is widespread in populations from western Canada. Overall genetic diversity of B. tectorum is much higher in eastern Canada than in the eastern U.S., while the genetic diversity in populations in western North America is similar between Canada and the U.S. The distribution of genetic diversity across Canada strongly suggests multiple introduction events. Heterozygous individuals, which are exceedingly rare in B. tectorum, were detected in three Canadian populations. Formation of novel genotypes through occasional outcrossing events could spark adaptive evolution and further range expansion across Canada of this exceedingly damaging grass.     

Phytochrome regulation of seed germination

Seed germination of many plant species is influenced by light. Of the various photoreceptor systems, phytochrome plays an especially important role in seed germination. The existence of at least five phytochrome genes has led to the proposal that different members of the family have different roles in the photoregulation of seed germination. Physiological analysis of seed germination ofArabidopsis thaliana (L.) Heynh. with phytochrome-deficient mutants showed for the first time that phytochrome A and phytochrome B modulate the timing of seed germination in distinct actions. Phytochrome A photo-irreversibly triggers the photoinduction of seed germination after irradiation with extremely low fluence light in a wide range of wavelengths, from UV-A, to visible, to far-red. In contrast, phytochrome B mediates the well-characterized photoreversible reaction, responding to red and far-red light of fluences four orders of magnitude higher than those to which PhyA responds. Wild plants, such asA. thaliana, survive under ground as dormant seeds for long periods, and the timing of seed germination is crucial for optimizing growth and reproduction. It therefore seems reasonable for plants to possess at least two different physiological systems for sensing the light environment over a wide spectral range with exquisite sensitivity of different phytochromes. This redundancy seems to enhance plant survival in a fluctuating environment.     

Population genetic structure of the seed pathogen Pyrenophora semeniperda on Bromus tectorum in western North America

We examined genetic variation in the ascomycete pathogen Pyrenophora semeniperda cultured from seeds of the invasive grass Bromus tectorum in the Intermountain West of North America. We sequenced the internal transcribed spacer (ITS) region of the nuclear ribosomal RNA genome in 417 monoconidial cultures collected from 20 sites in Washington, Idaho, Utah and Colorado, USA. ITS sequence diversity was surprisingly high; 12 unique haplotypes were identified, averaging 1.3% pairwise sequence divergence. All sites had at least two haplotypes present, and three sites had seven or more. One haplotype composed 60% of the isolates and occurred at all 20 locations; the remaining haplotypes generally occurred at low frequencies within sites but at multiple sites throughout the region. Sites in Washington and Idaho were more diverse than those in Utah and Colorado, averaging two more haplotypes and 67% more pairwise differences among haplotypes at a site. Analysis of molecular variance (AMOVA) indicated that more than 80% of the genetic variation was found within sampling locations, while 7-11% of the variation can be attributed to differences between northern (Washington and Idaho) and southern (Utah and Colorado) populations. The wide distribution of even uncommon haplotypes among sampling sites and weak correlations between genetic and geographic distances among populations (< 0.2) suggested that these populations recently were established from a common source. We hypothesize that the strains of P. semeniperda infecting B. tectorum in western North America probably arrived with the invasive grass from its native Eurasian range.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.     

绒毛番龙眼种子萌发生态特性的研究

就温度、光照、土壤水分条件对绒毛番龙眼 ( Pometia tom entosa( Bl.) Teysm.et Binn.)种子萌发的影响及种子寿命进行了研究 ,结果表明 ,绒毛番龙眼种子萌发的适宜温度为 2 0～ 35°C,最适温度为 30°C;周期性光照条件下的萌发优于全黑暗条件 ;适宜土壤含水量为 2 0 %～ 70 % ,最适为 6 0 % ;在室内自然摊放条件下 ,9d后发芽率减半 ,16 d后完全丧失发芽能力。研究认为 ,目前绒毛番龙眼的濒危状态主要是由于滥砍乱伐和森林破坏造成的 ,由于其种子具有顽拗性种子的一些特点 ,该物种宜采取活体保存的方法 ,以就地保护为主 ,活植物迁地保护为辅。     

Controls over invasion of Bromus tectorum: The importance of climate,soil, disturbance and seed availability

Question: Predicting the future abundance and distribution of invasive plants requires knowing how they respond to environmental conditions. In arid and semi‐arid ecosystems where water is a limiting resource, environmental conditions and disturbance patterns influence invasions by altering acquisition and utilization of water over space and time. We ask: 1. How do variations in climatic and soil properties influence temporal soil water dynamics? 2. How does this variation affect the establishment of Bromus tectorum (cheatgrass), a cool‐season annual grass that has successfully colonized much of the U.S. Great Basin? Location: Short‐grass Steppe in northeastern Colorado, USA; Arid Lands Ecology reserve in southeastern Washington, USA; and the Patagonian steppe of the Chubut province in Argentina. Methods: We utilized a soil water model to simulate seasonal soil water dynamics in multiple combinations of climatic and soil properties. In addition, we utilized a gap dynamics model to simulate the impact of disturbance regime and seed availability on competition between B. tectorum and native plants. Results: Our results suggest that climate is very important, but that soil properties do not significantly influence the probability of observing conditions suitable for B. tectorum establishment. Results of the plant competition model indicate that frequent disturbance causes more Bromus tectorum in invaded areas and higher seed availability causes faster invasion. Conclusions: These results imply a general framework for understanding Bromus tectorum invasion in which climatic conditions dictate which areas are susceptible to invasion, disturbance regime dictates the severity of invasion and seed availability dictates the speed of invasion.     

外来物种互花米草种子萌发的生态适应性

在3种条件即不同萌发温度和光照、不同贮藏温度和时间、不同盐浓度下,研究了互花米草种子萌发及胚生长特性.结果表明:在相同温度下,种子的萌发不受光照或黑暗条件影响,最适萌发温度为16/26℃(夜/昼)和25℃,萌发率高于90%;在-5℃、0℃、5℃、10℃的贮藏温度下,贮藏120 d的种子萌发率高于74%,表明种子能够安全越冬;当盐浓度为75～225 mmol·L~(-1)时,萌发率高于90%,150 mmol·L~(-1)的盐浓度较适宜胚的生长,胚芽鞘和胚轴的生长先于胚芽和胚根,盐溶液对胚轴、胚芽鞘的生长所产生的抑制作用小于胚根、胚芽,有利于已萌发的幼苗快速出土,适应多变的潮间带环境.