Seeds use temperature cues to ensure germination under nurse-plant shade in xeric Kalahari savannah

BACKGROUND AND AIMS: In arid environments many plant species are found associated with the canopies of woody perennials. Favourable conditions for establishment under canopies are likely to be associated with shade, but under canopies shade is distributed patchily and differs in quality. Diurnal temperature fluctuations and maximum temperatures could be reliable indicators of safe sites. Here, an examination is made as to whether canopy-associated species use temperature cues to germinate in shade patches, rather than matrix areas between trees. METHODS: The study was carried out in arid southern Kalahari savannah (Republic of South Africa). Perennial and annual species associated with Acacia erioloba trees and matrix species were germinated at temperature regimes resembling shaded and unshaded conditions. Soil temperature was measured in the field. KEY RESULTS: Germination of all fleshy-fruited perennial acacia-associated species and two annual acacia-associated species was inhibited by the temperature regime resembling unshaded conditions compared with at least one of the regimes resembling shaded conditions. Inhibition in perennials decreased with seed mass, probably reflecting that smaller seedlings are more vulnerable to drought. Germination of matrix species was not inhibited by the unshaded temperature regime and in several cases it increased germination compared with shaded temperature regimes or constant temperature. Using phylogenetically independent contrasts a significant positive relationship was found between canopy association and the germination at shade temperatures relative to unshaded temperatures. CONCLUSIONS: The data support the hypothesis that canopy species have developed mechanisms to prevent germination in open sun conditions. The results and data from the literature show that inhibition of germination at temperature regimes characteristic of open sun conditions can be found in fleshy-fruited species of widely divergent taxonomic groups. It is predicted that germination mechanisms to detect canopy shade based on temperature cues are widespread in species depending on nurse plants, especially bird-dispersed species.     

Effects of light and temperature on seed germination of cacti of Brazilian ecosystems

Environmental factors are used by plants as spatio‐temporal indicators of favorable conditions for seed germination. Thus, the objective of this study was to determine the effect of light and temperature on seed germination of 30 taxa of Cactaceae occurring in northeastern Brazil and to evaluate whether fluctuations in temperature are capable of altering light sensitivity. The seeds were tested for germination under two light conditions (12 h photoperiod and continuous darkness) and 10 temperature treatments: eight constant temperatures (10, 15, 20, 25, 30, 35, 40 and 45°C) and two alternating temperatures (30/20°C and 35/25°C). The species studied showed two photoblastic responses. All cacti from the Cactoideae subfamily (22 taxa) were classified as positive photoblastic (i.e., no germination in darkness), regardless of the temperature treatment used. Likewise, temperature fluctuation did not alter the seed sensitivity to light. On the other hand, the species of the Opuntioideae (five taxa) and Pereskioideae (three taxa) subfamilies are indifferent to light (i.e., germinated both in the presence and absence of light). The cacti from the areas of Caatinga and Cerrado showed an optimal germination temperature of 30°C, while the species from Atlantic Forest and Restinga areas showed an optimal germination temperature of 25°C.     

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.     

野豌豆属4种植物种子萌发的积温模型分析

以青藏高原野豌豆属窄叶野豌豆(Vicia angustifolia)、山野豌豆(V. amoena)、歪头菜(V. unijuga) 3种野生植物与一种当地栽培植物救荒野豌豆(箭筈豌豆) (V. sativa) ‘兰箭3号’种子为材料, 在5、10、15、20、25及30 ℃下进行萌发实验, 应用种子萌发的积温模型对上述4种植物萌发对温度的响应特征进行了比较分析。结果表明: 1)基于萌发速率(1/T_g)对种子萌发温度最低温T_b值的估计受萌发率(g)的影响较小; 与此不同, 除‘兰箭3号’种子外, 对萌发最高温T_c值的估计, 受到g的显著影响。 这表明种群内所有种子个体萌发的T_b值相对恒定, 但T_c值在有些物种中变异较大; 2)基于重复概率单位回归分析估计的种子萌发T_b值与基于萌发速率估计的值较为接近; 而由此方法估计的T_c值则与萌发率为50%时的估计值较为接近; 3)相比多年生豆科植物歪头菜和山野豌豆, 一年生豆科植物箭筈豌豆‘兰箭3号’与窄叶野豌豆具有相对较低的T_b与T_c值; 4)积温模型可准确地预测休眠破除后豆科植物种子在不同温度条件下的萌发进程。     

VARIATION IN GERMINATION RESPONSE TO TEMPERATURE IN RUBBER RABBITBRUSH (CHRYSOTHAMNUS NAUSEOSUS: ASTERACEAE) AND ITS ECOLOGICAL IMPLICATIONS

Seed collections from 72 rubber rabbitbrush populations occupying a range of habitats in western North America were incubated at 3 C in the laboratory. Collections from warm desert habitats required less than 2 weeks to achieve 90% relative germination under these conditions, while collections from montane habitats showed delayed germination requiring up to 20 weeks. When 13 representative collections were incubated at constant temperatures from 5 to 30 C, all germinated completely at 30 C within 4 weeks. Collections from warm desert habitats germinated rapidly over the whole range of temperatures. Montane collections sometimes exhibited dormancy at intermediate temperatures (15 and 25 C) even though they were ultimately able to germinate at lower temperatures. Results suggest that dormancy is conditional and temperature-dependent in this species. Chilling the seeds extends the temperature range for germination downward to include the chill temperature itself. Germination response to temperature and its variation as a function of habitat are of apparent adaptive significance, serving to time germination so that the probability of seedling survival is maximized in each habitat. Within populations, response to temperature varied as a function of year of harvest and of within-year harvest date, indicating that germination patterns are probably not under rigid genetic control but represent an integration of genetic and environmental factors.     

种子萌发的积温效应——以青藏高原东缘的12种菊科植物为例

温度是影响种子萌发的重要的环境因素之一。该文以青藏高原东缘的12种菊科植物为研究对象, 结合Logistic函数和积温公式, 通过非线性回归方法估算种子萌发的最低温度和积温, 研究了种子萌发对不同温度的响应。研究结果表明: (1)青藏高原东缘的12种菊科植物种子萌发的最低温度的平均值为0 ℃, 积温的平均值为94.5 ℃·d。与前人的研究相比, 该研究中萌发的最低温度较低, 积温较高, 这是该区域菊科植物长期适应青藏高原特殊的温度环境的结果; (2)种子萌发的最低温度与积温之间存在着显著的负相关关系(p = 0.04)。萌发最低温度较低的物种积温较高, 避免了种子在多变的温度环境下较早萌发所遇到的风险; (3)种子大小与积温之间存在着显著的正相关关系(p = 0.01)。在萌发最低温度差别不大的情况下, 与大种子相比, 小种子萌发所需的积温较低, 萌发较快, 在群落演替的早期占有优势。     

Effect of day length on germination of seeds collected in Alaska

Day length control can effectively limit seed germination to favorable seasons, but this phenomenon has been studied in relatively few wild plants. I tested species from interior Alaska for day length control of germination under controlled conditions, and I also monitored germination phenology in natural habitats. Unstratified and cold-stratified seeds were germinated on short (13 h) and long (22 h) day length and in the dark at constant and alternating temperatures. On long day length, unstratified Ledum decumbens and Saxifraga tricuspidata seeds germinated from 5 C to 20 C, but on short day length few or no seeds germinated at 5 C and 10 C and germination was reduced at higher temperatures. Unstratified seeds of Diapensia lapponica and Chamaedaphne calyculata germinated only at 15 C and 20 C on long day length, and short day length completely inhibited germination. Cold stratification widened the temperature range for germination on both long and short day lengths, but germination was still lower on short than long day length. Germination phenology in natural habitats was consistent with germination in controlled conditions. In these species, short day length and low temperatures interact to inhibit germination in the fall. After overwintering, seeds germinate in the spring at low temperatures and on long day lengths. The inhibitory effect of short day length is not important in the spring because day length is already long at snowmelt.     

TEMPERATURE EFFECTS ON SEED GERMINATION OF EAST ASIAN AND TERTIARY RELICT SPECIES OF DIOSCOREA (DIOSCOREACEAE)

To obtain information relevant to the evolution of dormancy, germination responses to temperature of intact seeds and naked zygotic embryos were compared among taxonomically closely related species of Dioscorea in the section Stenophora. We examined five species from the northern half of the East Asian distribution area and four species considered to be Tertiary relict species from the Appalachians, the Caucasus, and the Balkans. Although features of germination of the intact seeds differed from species to species, the naked embryos of all species germinated over a similar wide range of temperatures without any marked differences among species. This unitary physiological feature of the embryos suggests the possibility that the temperature responses of the embryos have not changed since the Tertiary period. In the East Asian species, as the distribution area shifts to the north the seeds gradually lose their dormant features, and consequently the germination behavior of the seeds gradually changes to resemble that of their embryos. The seed of the northernmost species has no dormant features at all, and temperature responses of the seed are the same as those of the embryo. Full germination of the intact seeds of East Asian and Tertiary relict species required prior chilling treatment. Unlike the East Asian species, however, the relict species germinated to some extent at higher temperatures over a narrow range without prior chilling. However, the resultant germlings died or elongated poorly. Thus, the germination process of relict species may have become less sensitive to high-temperature inhibition after their isolation from Asian species.     

The effect of Sea Water and Temperature on the Germination Behaviour of Crithmum maritimum

The seeds of Crithmmm maritimum L. were germinated floating on various concentrations of sea water up to 50% at constant temperatures of 5, 10, 15, 20, and 25°C and at alternating temperatures of 5 and 15°C. 5 and 25°C. and 15 and 25°C. Significantly higher germination was obtained at alternating than at constant temperature. When two constant temperatures at which no germination occurred were alternated, good germination was obtained. There was reduced germination and increase in time of first germination as sea water concentration increased, in the absence of sea water, high temperature caused not only severe inhibition of germination but also permanent injury to the seeds. The results help to explain the germination behaviour of the species in nature.     

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.     

Strict requirement of fluctuating temperatures as a reliable gap signal in <Emphasis Type="Italic">Picris hieracioides</Emphasis> var. <Emphasis Type="Italic">japonica</Emphasis> seed germination

Picris hieracioides var. japonica (Asteraceae), which grows in occasionally disturbed habitats such as riverbanks, is rarely observed under dense vegetation. We examined the effect of the experience and timing of receiving leaf-transmitted light in gap-detecting seed germination in this plant. Seeds under unfiltered light, which simulated the light conditions of seeds on the soil surface in a canopy gap, germinated at a constant temperature of 20°C. However, most seeds in darkness, which simulated the light conditions of seeds buried in the soil without receiving leaf-transmitted light, germinated under temperature fluctuations of over 4°C. Seeds in darkness after receiving leaf-transmitted light for 1 week, which simulated the light conditions of seeds buried in the soil after receiving leaf-transmitted light, germinated under temperature fluctuations of over 8°C. Finally, seeds under continuous leaf-transmitted light, which simulated the light conditions of seeds on the soil surface below preexisting vegetation, germinated under temperature fluctuations of over 12°C. Seeds that experience unfiltered light, which suggests that they are in a gap, should not delay germination. In contrast, seeds that have received leaf-transmitted light should delay germination until the vegetation above is removed. Seeds exposed to leaf-transmitted light required larger temperature fluctuations in darkness than did untreated seeds, and seeds under continuous leaf-transmitted light required the largest temperature fluctuations. The various germination reactions to each gap signal in P. hieracioides var. japonica seeds allow the more reliable detection of gaps for subsequent seedling establishment. The requirement for gap signals that created high precision of timing in the germination process results in the germination of this species only in gaps. Therefore, P. hieracioides var. japonica is rarely found under dense vegetation.     

Seed germination of Pilosocereus arrabidae (Cactaceae) from a semiarid region of south‐east Brazil

We evaluated the effect of temperature regimes (six constant and four alternating temperatures), light qualities (five red : far red ratios) and water potentials (ΨW; seven NaCl and polyethylene glycol 6000 [PEG] solutions) on the percentage and germination rate, as well as the post‐seminal development morphology, that allow Pilosocereus arrabidae seeds to germinate in a hot semiarid climate on the south‐eastern Brazilian coast. The results showed that seeds germinated similarly between constant and alternating temperatures, with an optimal germination at 25/20°C and 20°C. Pilosocereus arrabidae seeds were photoblastic positive and the final germination percentage was inhibited at low red : far red ratios. Maximum germination was obtained in distilled water (0 MPa) and decreases of Ψ_W in the solutions reduced the germination, which was lower in NaCl than in iso‐osmotic PEG solutions. Germination inhibition appears to be osmotic because the recovery response was high when non‐germinated seeds from both iso‐osmotic solutions were transferred to water. Seeds of P. arrabidae are small and germination is phaneroepigeal. Despite the slow growth typically seen in seedlings and adults of Cactaceae, germination in this species depends on the ability of the seeds to appropriately sense and react to environmental cues that correlate with times and places under low‐risk growth conditions.     

Effect of temperature on seed germination of ten species and varieties ofLimnanthes

Ten accessions of seed of the genus Limnanthes, representing seven species and their varieties, were tested for germination at temperatures of 40°, 50°, 60°, 70° F, and at room temperature varying from 72°–78° F. Four of the ten accessions germinated best at 40°, three at 50°, and three at 60° F. Germination was relatively poor at 70° F and almost zero at room temperature. The total germination averaged over all accessions was about the same for both 40° F and 50° F, slightly less for 60° F. Seeds at 60° F reached a peak in germination rate in four days; those at 50° F required six days; those at 40° F required 11 days. Seeds, started at room temperature and moved to 50° F 15 days later, attained only about half the total germination of a similar group of seeds originlly started at 50° F. Some of the species tested germinated almost equally well at 40°, 50°, or 60° F; others germinated best only at, or near, one of these temperatures. Substantially all of the seeds that germinated over a period of 50 days did so in less than 30 days; most of these germinated in the first 14 days.     

The temperature dimension of the seed germination niche in fen wetlands

The seed germination niche partly determines adaptation, ecological breadth and geographic range in plant species. In temperate wetlands, environmental temperature is the chief regulator of germination timing, but the ecological significance of high and low temperatures during dormancy break and germination is still poorly understood. Our aim was to characterize the temperature dimension of the germination niche in mountain base-rich fens, determining (1) the effect of different temperatures on dormancy break and germination, and (2) whether different germination strategies may be identified at the species level. We conducted laboratory germination experiments with seeds of 15 species from these habitats, collected in 18 fen sites in the Cantabrian Mountains (Spain) for two consecutive years. In all the species, the seeds were totally or conditionally dormant at dispersal and stratification produced a significant increase of germination. In most cases, there was not an obligatory requirement for cold temperatures during dormancy break, since warm stratification promoted germination as well. Although the optimal germination thermoperiod was generally high (30/20 °C), most species could also germinate at lower temperatures after cold-stratification. We also identified a group of species associated to cold-water springs that germinated only at low temperatures. Our results demonstrate that dormancy break in mountain base-rich fens does not obligatorily depend on cold temperatures during overwintering. Furthermore, germination at cool temperatures may be more widespread in wetland habitats than previously thought. The existence of two distinctive germination strategies, ‘warm’ and ‘cool’, can potentially give rise to divergent species responses to climate change.     

Effects of extreme temperature on seedling establishment of nonnative invasive plants

Climate change and plant invasion are two of the most important ecological issues facing the world today. Extreme events are likely to play an important role in plant invasion. For example, tolerance to temperature stress is critical for plant germination and survival of seedlings. Nonnative invasive species tend to differ from co-occurring native species in several traits. Increased mean temperatures are known to enhance the risk of plant invasions, but few experimental studies have linked plant invasion to both increasing mean temperature and extreme (low and high) temperatures. Ten plant species from Asteraceae (six nonnative invasive and four native species) were chosen and six temperatures (extremely low, average winter, average annual, average summer, high and extremely high) were used to test the effects of extreme temperatures on plant invasion in southern China. The results showed that nonnative invasive plant species (IS) germinated more readily and the seedlings grew better than those of native plant species (NS) at high temperatures, suggesting that global warming may facilitate invasion. Extreme temperatures decreased the seed germination rate and seedling growth of both IS and NS, although NS were more tolerant of extremely low temperatures (5/0 °C). IS, in turn, were more tolerant of extremely high temperatures (40/35 °C). Extreme high temperatures may increase the risk of plant invasion because IS seedlings are better able to become established, whereas low temperatures may hinder invasion. In addition, the species-specific differences in plant origin (IS and NS) and temperature tolerance were correlated with other climatic factors and should be considered in managing invasive species in a changing world.     

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.     

Seed‐based approach for identifying flora at risk from climate warming

In obligate seeding species, the germination niche is crucial for colonization and population survival. It is a high‐risk phase in a plant's life cycle, and is directly regulated by temperature. Seeds germinate over a range of temperatures within which there is an optimum temperature, with thresholds above and below which no germination occurs. We suggest that abrupt changes in temperature associated with a warming climate may cause a disconnect between temperatures seeds experience and temperatures over which germination is able to occur, rendering obligate seeding species vulnerable to decline and extinction. Using a bidirectional temperature gradient system, we examined the thermal constraints in the germination niche of some geographically restricted species from the low altitude mountains of the Stirling Range, southern Western Australia, including seedlots from lowland populations of four of these species. We demonstrated that high temperatures are not a limiting factor for germination in some restricted species, signifying a lack of relationship between geographic range size and breadth of the germination niche. In contrast, we identified other restricted species, in particular Sphenotoma drummondii, as being at risk of recruitment failure as a consequence of warming: seeds of this species showed a strong negative relationship between percentage germination and increasing temperature above a relatively low optimum constant temperature (13°C). We found some ecotypic differences in the temperature profiles between seeds collected from montane or lowland populations of Andersonia echinocephala, and while specific populations may become more restricted, they are perhaps at less risk of extinction from climate warming. This seed‐based approach for identifying extinction risk will contribute tangibly to efforts to predict plant responses to environmental change and will assist in prioritizing species for management actions, directing limited resources towards further investigations and can supplement bioclimatic modelling.     

Spatial relationships between cacti and nurse shrubs in a semi-arid environment in central Mexico

Abstract. The Zapotitlán de las Salinas valley, central Mexico, harbours a high diversity of cacti. Pattern analysis indicated that the establishment of two columnar cacti, Neobuxbaumia tetetzo and Cephalocereus hoppenstedtii, and of three small globose cacti, Coryphantapallida, Mammillaria colina andM casoi, is aggregated and associated with perennial nurse shrubs. Some nurse species, Castela tortuosa, Caesalpinia melanadenia and Eupatorium spinosarum have a higher number of cacti beneath their canopies than would be expected by chance. A replacement pattern was found between the columnar cacti and their nurses, an aspect which was not found with the globose cacti. Following the assumption that protection against excessive radiation is the main factor determining the nurse effect, the azimuth orientation of the cacti with respect to their nurses was evaluated. Only Coryphanta pallida presented a non-random distribution with a tendency towards the North and West. The difference in maximum temperature between the soil surface under the different nurse species and of open spaces, which is reached at midday, was 16 °C. No significant differences were found in beneath-canopy temperatures for the three nurse species considered. Soil nitrogen levels were significantly lower beneath the different nurse plants than in open spaces. This result suggests that soil fertility is not an important factor in the nurse-plant phenomenon in Zapotitlán.     

The Effects of Light and Temperature on Germination and Growth of Luffa aegyptiaca

The effects of light and temperature on the germination and growth of Luffa aegyptiaca were investigated both in the laboratory and in the field. The seeds germinated in both darkness and light but germination was better in the light. At constant temperatures germination was best at 21°C, while alternating temperatures of 21 and 31°C and 15 and 41°C caused higher germination than the most favourable constant temperature. Constant temperatures of 15 and 31°C and alternating temperatures of 21 and 41°C resulted in very low germination, whereas no germination occurred at 41°C and at alternating temperatures of 31 and 41°C. Soil depth caused only a delay in seed germination, as it did not affect the total germination. High temperature and high light intensity resulted in good seedling growth in terms of dry weight, leaf area and relative growth rate. High temperature and low light intensity caused increased plant height and high shoot weight ratio, both of which manifested in seedling etiolation. They also caused high leaf area ratio. Under low temperatures, irrespective of light intensity, growth was generally poor, but it was significantly poorer under low light intensity, which also caused high root weight ratio. High light intensity was principally responsible for high leaf weight ratio. The results help to explain the abundance of the species in newly cleared areas in Lagos and its environs.