Seedling tolerance to cotyledon removal varies with seed size: A case of five legume species

It is generally accepted that seedlings from large seeds are more tolerant to defoliation than those from small seeds due to the additional metabolic reserves present in the large seeds. However, information on the effects of amount of seed reserves (cotyledon removal) from seedlings resulting from large vs. small seeds on seedling growth and long‐term survival in the field is limited. Five legume species with different sizes of seeds were sown in the field and none, one, or both cotyledons removed 7 days after seedling emergence. Seedling biomass, relative growth rate (RGR) and survival were determined at different time. Cotyledon removal, species, and their interaction had significant effects on seedling growth and survival. During the period between 33 and 70 days, seedlings from large seeds had a significantly lower RGR than those from small seeds. Biomass, RGR, and survival of seedlings from large seeds were significantly reduced by removal one or both cotyledons, whereas those of seedlings from small seeds were not affected. Seed energy reserves are more important for the early growth of seedlings from large seeds than for those from small seeds. The overall effect of cotyledon removal on growth and survival varies with seed size (i.e., energy reserves) with seedlings from small seeds being less sensitive than those from large seeds under field conditions.     

Seed size, seedling morphology, and response to deep shade and damage in neotropical rain forest trees

To investigate the existence of coordinated sets of seedling traits adapted to contrasting establishment conditions, we examined evolutionary convergence in seedling traits for 299 French Guianan woody plant species and the stress response in a shadehouse of species representing seed size gradients within five major cotyledon morphology types. The French Guianan woody plant community has larger seeds than other tropical forest communities and the largest proportion of hypogeal cotyledon type (59.2%) reported for tropical forests. Yet the community includes many species with intermediate size seeds that produce seedlings with different cotyledonal morphologies. A split-plot factorial design with two light levels (0.8% and 16.1% PAR) and four damage treatments (control, seed damage, leaf damage, stem damage) was used in the shadehouse experiment. Although larger-seeded species had higher survival and slower growth, these patterns were better explained by cotyledon type than by seed mass. Even larger-seeded species with foliar cotyledons grew faster than species with reserve-type cotyledons, and survival after stem grazing was five times higher in seedlings with hypogeal cotyledons than with epigeal cotyledons. Thus, to predict seedling performance using seed size, seedling morphology must also be considered.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

Seed reserve translocation and early seedling growth of eight tree species in a tropical deciduous forest in Mexico

Seed reserves play an essential role during germination and seedling establishment and are particularly important for species that grow in seasonal ecosystems with a short growing season. In this study, we examined (a) how and when the seedlings change their dependence from seed resources to external resources, (b) the lipid, nitrogen, and non-structural carbohydrate reserve translocation from seeds to seedlings over time, and (c) whether reserve translocation may be correlated to cotyledon and leaf lifespan of seedlings for eight tree species in a tropical deciduous forest in north-western Mexico. Our results showed that the cotyledon lifespan was not related to the cotyledon type (photosynthetic or reserve) and that the cotyledon biomass did not decrease significantly until germination. In six of the eight studied species, biomass allocation to the leaves was favored; lipids were the first reserve exhausted before the first leaves were totally expanded in seven of the eight study species. Species with the highest N concentration had expanded leaves and lost their cotyledons faster than species with a low N concentration. Our results suggest that tropical deciduous forest species employ different strategies to survive the dry season and re-sprout in the next growing season mediated by seed reserve concentrations, translocation patterns and subsequent biomass allocation.     

Impacts of secondary seed dispersal and herbivory on seedling survival in Aesculus turbinata

Abstract. Aesculus turbinata is a tree species with large seeds (6.2 g mean dry weight). We studied the demography of its seeds and seedlings in a temperate deciduous forest in northern Japan to elucidate the ecological significance of large seeds with special reference to herbivory and secondary dispersal. Both seed and seedling stages suffered greatly from herbivores. Seedling herbivory was important judged from experiments with shoot clipping and hypogeal cotyledon removal. However, some seedlings survived through re-sprouting after herbivory. Survival rate and percentage resprouting seedlings were lower than those with remaining cotyledons, though seedling size was not affected. This suggests that stored resources in hypogeal cotyledons are working as a kind of ‘risk hedge’ against severe aboveground shoot clipping experienced by A. turbinata. The spatial distribution of seedlings was expanded via seed scatter-hoarding by rodents. Seedling survival rate was higher within canopy gaps than under closed canopy, indicating that canopy gaps are safe sites for establishment, and was negatively correlated with seedling density. Therefore, secondary seed dispersal in this species seems to be effective in ‘finding’ safe sites and in ‘escaping’ density-dependent mortality. The large seeds and seedlings of A. turbinata are attractive to herbivores, but the high resistance of seedlings to herbivory due to large reserves and the effective secondary dispersal appear to mitigate these disadvantages.     

The Influence of Sowing Depth and Seed Size on Seedling Emergence Time and Relative Growth Rate in Spring Cabbage (Brassica oleracea var. capitata L.)

The effects of seed size and sowing depth on the time of seedlingemergence and on the growth of spring cabbage studied in greenhouseand field experiments. Seed size had little effect on seedlingemergence time, but plants derived from large seeds were largerthan those from small seeds. Increased sowing depth delayedseedling emergence and reduced seedling relative growth rate(RGR). To our knowledge, these effects of sowing depth on RGRindependent of inter-plant competition have not been reportedpreviously for any species. Sowing depth had no effect on thenet assimilation rate (NAR) of seedlings, indicating that thelower RGR of seedlings from deep sowings was associated witha low light interception by small cotyledons which in turn resultedfrom disproportionately low partitioning of assimilates to thecotyledons during pre-emergence growth in favour of producinghypocotyls of greater length. Brassica oleracea, specific leaf area, growth analysis, dry matter partitioning, light interception, cabbage, seed size, sowing depth, seedling emergence time, relative growth rate, net assimilation rate     

Growth and survival of cork oak (Quercus suber) seedlings after simulated partial cotyledon consumption under different soil nutrient contents

Aims

We examined the importance of partial seed consumption (cotyledon loss) by rabbits in the early establishment of seedlings of cork oaks restricted to nutrient-impoverished soils.

Methods

To determine the importance of cotyledons in the growth and development of seedlings, we simulated two levels of predation [light (30 % cotyledon loss) and heavy (60 % loss) partial consumption] and two soil nutrient contents (nutrient-poor soil, nutrient-rich soil). Seedlings height, root length, dry root and shoot biomass, specific leaf mass, leaf density, gas exchange, chlorophyll fluorescence parameters and photosynthetic pigment concentrations were determined.

Results

Results indicated that effect of nutrient level on the growth of the oak seedlings was more important than that of cotyledon biomass. However, in nutrient–poor soils, cotyledon biomass played a major role in the early performance of cork oaks. Acorns grown in nutrient-rich substrate, despite having greater aerial vigor, were slower to develop a vertical root, and hence less likely to reach permanent moisture. Cotyledon loss caused a decrease in the biomass of roots and shoots when acorns were heavily consumed, and as a result experienced a reduction in net photosynthetic rate, stomatal conductance and chlorophyll concentration. Survival of seedlings was unaffected by either soil type or cotyledon loss.

Conclusions

Our results show that effects of soil type on the survival of oak seedlings were more important than those of cotyledon biomass. However, in a competitive situation, cotyledon biomass, as an indicative of growth nutrient support rather than an energy source, could be vital in a nutrient-poor environment, particularly in Mediterranean climate regions and for species with little inherent drought tolerance (as is the case of Quercus spp.), where rapid root growth is required to ensure that contact with soil moisture is maintained over the first summer.     

An Advantage of Large Seed Size: Tolerating Rather than Succumbing to Seed Predators1

Although large seeds might be more attractive and apparent to seed predators, large seed size could enable tolerance of seed predators. If seeds are large enough to sustain damage that would kill smaller seeds yet still produce viable seedlings, investment above the minimum by the maternal plant could be advantageous. I tested this hypothesis by removing 0–80 percent of the cotyledons of four large-seeded (4–180 g) tree species from Papua New Guinea and monitoring germination and seedling growth for eight months. All species showed little negative effect on seedling size with up to 50 percent removal of cotyledons and the larger species showed a less serious effect on growth than smaller-seeded species above 50 percent removal. Large-seeded species clearly have more than the minimum-required cotyledonary reserves. Observations of viable seedlings with heavily damaged cotyledons suggest that these species withstand attack by rodents and beetles by virtue of their large size.     

南亚热带森林24种乔木的种子萌发和幼苗生长

以膨胀珍珠岩为基质,在光和暗的条件下,对２４种南亚热带森林乔木的种子萌发和幼苗生长进行了研究。种子的形态和重量与种的演替阶段有关,种子的重量也与不同的种有关。不同种的萌发率差异较大,种子较大的种,萌发率较高。肉质果实的种子,开始萌发的时间较长。光和暗条件对萌发率和萌发速度无明显影响。幼苗高度和种子重量呈正相关,这一相关在光条件下比暗条件下更明显。暗条件明显地引起群落演替早期的树种的茎徒长,限制根的生长,而对演替后期种无显著的影响。幼苗地上部分和根系生物量的分配与种的演替阶段有关,也与不同的科有关。幼苗地上部分和根系生物量的分配也受光暗条件的影响,而不受种子重量的影响。根和叶的生物量分配和种在其群落演替阶段有关。幼苗的生物量与种子重量有显著的正相关。幼苗的相对生长率相差较大,在没有营养供应的生长基质中,以演替中间阶段的种的相对生长率较大。幼苗从种子的物质利用效率与种子重量呈负相关,而与种的演替阶段无明显的相关。     

辽东栎幼苗生长对种子大小和子叶去除处理的响应

该文在温室内盆播条件下,设置不去除子叶(对照)、去除1/4子叶(轻度子叶去除)、1/2子叶(中度子叶去除)和3/4子叶(重度子叶去除) 4个处理,研究了种子大小和子叶去除处理对辽东栎幼苗生长及其形态学可塑性的影响。结果表明:(1)大种子幼苗株高、基茎、叶片数、单株叶面积、总干质量和根冠比在所有处理均大于小种子幼苗,除株高外,上述其他参数在轻度子叶去除处理的不同大小种子幼苗间均差异显著(P0.05),前者总干质量在中度子叶去除处理与后者差异也显著(P0.05)。大种子幼苗比叶面积和比根长在各处理均大于小种子幼苗,比枝长则相反。(2)随着子叶去除强度增大,大、小种子幼苗株高、基茎、叶片数、单株叶面积和总干质量等均呈减小趋势,株高、基径、叶片数和单株叶面积在重度子叶去除处理均显著小于对照和轻度子叶去除处理;小种子幼苗对子叶损失反应更为敏感,其株高、叶片数、单株叶面积和总干质量在中度子叶去除处理也显著小于对照(P0.05);小种子幼苗根冠比随子叶去除强度增大而增大,在中度和重度子叶去除处理显著大于对照和轻度子叶去除处理(P0.05)。(3)随着子叶去除强度增大,大小种子幼苗的比叶面积和比根长均逐渐增大,但大种子幼苗比枝长持续减小,轻度子叶去除处理的小种子幼苗比枝长最大。     

Host compatibility of the cloud forest mistletoe Psittacanthus schiedeanus (Loranthaceae) in Central Veracruz, Mexico

The consequences of the seed deposition of the parasitic mistletoe Psittacanthus schiedeanus were evaluated in a 32-mo study. We conducted a field seed inoculation experiment to determine variation in seed adhesion, seed germination, seedling establishment, and plant survival to reproduction among five host species and to evaluate whether these post-dispersal processes explain mistletoe prevalence and specificity at the regional scale. Seeds without an exocarp were inoculated onto branches of individuals of the five most common host species identified in nature in central Veracruz, México. Seed fate was monitored for 2 yr, at weekly intervals for the first 2 mo and at 2-mo intervals thereafter. The height and diameter of experimental host branches and canopy cover above them were measured to see if these factors affected mistletoe establishment. Significant differences in seed attachment and seed germination were found among host species. Fewer seeds remained attached on experimental branches of Quercus germana than those of Liquidambar styraciflua, Acacia pennatula, and Platanus mexicana. Although significant differences in seed germination were observed among species (significantly greater on A. pennatula), >70% of mistletoe seeds germinated within the first 5 wk on all host species. Towards the end of the inoculation experiment, more mistletoe seedlings survived, grew, and then flowered on Liquidambar styraciflua than on A. pennatula, P. mexicana, Q. germana, or Q. leiophylla. Host branch initial height and diameter did not affect seedling survival, but seedlings survived better on trees where the canopy was more open. Our results suggest that Liquidambar styraciflua is the most compatible host species with P. schiedeanus in central Veracruz. Not surprisingly, Liquidambar is by far the most common host tree for P. schiedeanus in this area as well. We suggest that the observed local specialization is a result of seed dispersal as consequence of bird foraging and territorial behavior, host abundance, and host species compatibility.     

Impact of Cotyledon and Leaf Removal on Seedling Survival in Three Tree Species with Contrasting Cotyledon Functions1

The relative importance of cotyledons and leaves for seedling survival was evaluated using a factorial field experiment on three neotropical tree species with contrasting cotyledon functional morphologies (photosynthetic, epigeal reserve vs. hypogeal reserve). In all species, cotyledon and leaf removal shortly after leaf expansion had additive negative effects on seedling survival over 7 weeks. Carbon supplies from cotyledons and other carbohydrate reserves apparently enhanced ability of seedlings to cope with herbivory and disease.     

Canopy arthropods community within and among oak species in central Mexico

Quercus rugosa and Q.laurina are species that presents a wide geographical distribution range in temperate forests of Mexico. Oak canopies contain a considerable portion of arthropod diversity and the arthropods fauna fulfill a wide variety of ecological roles. We examined the effect of oak species and seasonal changes on some community structure parameters (diversity, composition, similarity, biomass, rare species, and density of arthropod fauna) of canopy arthropods. In total, 40 oak canopies were fogged during rainy and dry season. A total of 614 identified arthropod morphospecies were recognized belonging to 22 orders associated with tree canopies. A separation of host tree species during both seasons, suggesting a different community structure on host plants species was demonstrated by the principal component analyses (PCA), therefore, differences between oak species results in phenotypes that structure the composition of the arthropod community. Q.laurina registered the highest densities, diversity index and number of rare species in comparison with Q.rugosa. While arthropod biomass showed an inverse pattern. Trees more close to one another (spatial distance) register a more similar canopy arthropod fauna. This study suggests that the trees of Q.laurina could act as a center of biodiversity by the accumulation of arthropod fauna with a considerable number of rare species, which presents wide ecological roles or is involved in critical processes that maintain forest ecosystems[Current Zoology 55(2):132-144,2009].     

Old-field seedling responses to insecticide,seed addition,and competition

The controls of seedling emergence and survival determine the potential distribution of adult plants and, thereby, plant community structure. Seed availability, competition from established neighbors, and seedling predation may all limit seedling recruitment. In this field experiment, we followed the emergence and survival of seedlings of three perennial forbs, Achillea millefolium, Hypericum perforatum, and Monarda fistulosa, in old-fields in southeastern Michigan, USA. As adults, all three have aromatic foliage that may deter herbivory, but seedlings may be more susceptible than adults. To establish the relative importance of potential controls on seedling numbers, we manipulated seed availability through seed additions, the influence of competitors by neighbor-removals, and the influence of insect herbivores with insecticide in a fully factorial field experiment. Seed addition and insecticide never affected seedling emergence for any species. Competition from established neighbors controlled seedling emergence for all three species and decreased Achillea survival. Insecticide significantly increased Monarda seedling survival in competition plots, significantly increased Hypericum survival in open plots, and had no effect on Achillea. Notably, insecticide increased survival of the native Monarda fistulosa more than the two introduced species. While neighbors strongly reduced emergence and survival of all three species, herbivores acted on a species-specific basis. These results suggest the differential effects of insects may contribute more to the seedling species composition and abundance patterns than the less-selective influence of competition.     

Seed size variation in the palm Euterpe edulis and the effects of seed predators on germination and seedling survival

Intraspecific variation in seed size is common in wild plant populations and has important consequences for the reproductive success of individual plants. Multiple, often conflicting evolutionary forces mediated by biotic as well as abiotic agents may maintain such a variation. In this paper we assessed seed size variation in a population of the threatened, commercially important palm Euterpe edulis in southeast Brazil. We investigated (i) how this variation affects the probability of attack by vertebrate and invertebrate post-dispersal seed predators, and (ii) if seed size influences the outcome of seeds damaged by beetles in terms of seed germination and early survival of seedlings. Euterpe edulis seeds varied in diameter from 8.3 to 14.1 mm. Neither insects nor rodents selected the seeds they preyed upon based on seed size. Seed germination and total, shoot and root biomasses of one-year seedlings were significantly and positively affected by seed size. Root biomass and seedling survival were negatively affected by seed damage caused by a scolytid beetle (Coccotrypes palmarum) whose adults bore into seeds to consume part of the endosperm, but do not oviposit on them. Seed size had a marginally significant effect on seedling survival. Therefore, if any advantage is accrued by E. edulis individuals producing large seeds, this is because of greater seed germination success and seedling vigor. If this is so, even a relatively narrow range of variation in seed size as observed in the E. edulis population studied may translate into differential success of individual plants.     

Seed-mass effects in four Mediterranean Quercus species (Fagaceae) growing in contrasting light environments

Three hypotheses have been proposed to explain the functional relationship between seed mass and seedling performance: the reserve effect (larger seeds retain a larger proportion of reserves after germinating), the metabolic effect (seedlings from larger seeds have slower relative growth rates), and the seedling-size effect (larger seeds produce larger seedlings). We tested these hypotheses by growing four Mediterranean Quercus species under different light conditions (3, 27, and 100% of available radiation). We found evidence for two of the three hypotheses, but none of the four species complied with all three hypotheses at the same time. The reserve effect was not found in any species, the metabolic effect was found in three species (Q. ilex, Q. pyrenaica, and Q. suber), and the seedling-size effect in all species. Light availability significantly affected the relationships between seed size and seedling traits. For Q. ilex and Q. canariensis, a seedling-size effect was found under all three light conditions, but only under the lowest light (3%) for Q. suber and Q. pyrenaica. In all species, the correlation between seed mass and seedling mass increased with a decrease in light, suggesting that seedlings growing in low light depend more upon their seed reserves. A causal model integrates the three hypotheses, suggesting that larger seeds generally produced larger seedlings.     

Seed mass versus seedling performance in Scots pine: a maternally dependent trait

It is generally accepted that larger seeds give rise to seedlings with better performance. On the other hand, the size that a seed reaches is genetically determined by at least two different traits ; the genetic variability of the developing embryo and the genetic variability of the maternal plant. Thus, the relative contributions of these two traits affect seedling performance by influencing seed size. In this paper, I investigate the effect of seed size on seedling performance in the Scots pine ( Pinus sylvestris ). From eight maternal plants, 50 seeds were planted in each of two soil types (800 seeds in total), and seedling performance was monitored for 1 yr. Seed mass proved to be highly constant within maternal plants. Soil type influenced emergence and survival; however, the effect of soil type differed depending on maternal origin. Seed mass was positively correlated with seedling emergence, although this relationship was not found for seedling survival or date of emergence. The initial growth of the shoot was also positively correlated with seed mass. However, after one growing season, seed mass had no effect on seedling performance, which depended exclusively on maternal origin. Nevertheless, the mean mass of seeds produced by plants was positively correlated with mean values of growth parameters. Thus, first-year seedling performance seems to be a maternal trait indirectly associated with seed size.     

Additive and nonadditive effects of herbivory and competition on tree seedling mortality, growth, and allocation

The interaction between simulated cotyledon herbivory and interspecific competition was studied in a greenhouse experiment using two species of trees, Acer rubrum and Quercus palustris, which commonly invade abandoned agricultural fields. Herbivory treatments were applied as a gradient of cotyledon removal for A. rubrum with 0, 25, 50, 75, and 100% of cotyledon tissue removed. Cotyledons from Q. palustris were clipped and removed (control, early, and late removal) to create a gradient of seed reserve availability. The competition treatment consisted of plugs of old-field vegetation that filled the pots with perennial cover. Mortality of seedlings was higher with competition. There was a significant interaction between herbivory and competition with the highest mortality occurring with competition at the highest intensity of herbivory in both species. Herbivory reduced biomass for Q. palustris only, while competition reduced biomass in both species. Neither species showed an interaction between herbivory and competition for growth. There was a significant interaction between herbivory and competition on allocation patterns for both species, with greater allocation to roots with competition at the highest intensity of herbivory. This study demonstrates the potential for cotyledon herbivory and competition to interact, altering the invasion of tree seedlings into abandoned agricultural land.     

Effects of Planting Depth,Sediment Grain Size,and Nutrients on Ruppia maritima and Potamogeton perfoliatus Seedling Emergence and Growth

Protocols are now available for seed harvest, storage and germination of several mesohaline and polyhaline species; however, low seedling survival rates point to the need for an increased understanding of factors affecting seedling establishment. Depth of seed burial in sediments and initial seedling growth rates are shown to be limiting factors for photosynthetic competency of Ruppia maritima and Potamogeton perfoliatus. Seedling emergence is inversely proportional to planting depth on sediments ranging in grain size from coarse sands (850 μm) to silt (63 μm). Less than 6% of the seeds of either species emerged when buried to a depth of 3 cm in test sediments. Germination was greatest for seeds placed on the surface of sediments; however, these seedlings were subject to displacement because of the weak and fragile roots produced during early growth. Fine sediments may be more favorable for R. maritima seedling establishment, because seedling emergence and height decreased with increasing sediment grain size. Potamogeton perfoliatus seedlings seem to be more tolerant of a wider range of sediment grain sizes than R. maritima as indicated by the lack of an effect of sediment grain size on P. perfoliatus seed emergence, seedling height, and biomass. Increasing nutrients stimulated seedlings of both species; however, even at the highest concentrations tested, growth, as determined by shoot elongation and leaf and root formation, slowed within 7–10 days. This suggests factors other than mineral nutrients and light limit growth or that growth shifts from aboveground biomass production to belowground vegetative spread.     

杉木种子萌发及幼苗生长对光强的响应

设置不同光强梯度(透光率分别为100%、40%、20%、10%和5%,光照强度PPFD分别为201.3、77.0、37.5、19.2、9.8 μmol·m^-2·s^-1),研究光对杉木种子萌发和幼苗早期生长的影响,分析杉木种子萌发、幼苗存活、生长、形态响应、生物量积累及其分配格局对不同光环境的响应策略.结果表明: 杉木种子的萌发率、存活率、建植率和萌发指数在不同光强梯度下均有显著差异,且40%的透光率是种子最适萌发条件,萌发率最高,而全光照下存活率和建植率最高;随光照强度的减弱,杉木幼苗茎长增大,根长、子叶长、子叶厚、真叶数呈降低趋势,而基径在各光照强度间无显著差异;总生物量、根生物量、茎生物量、叶生物量均表现为全光照下最大.随着光照强度的减弱,光合组织与非光合组织生物量比、叶生物量比呈降低趋势,茎生物量比呈增加趋势,根冠比和根生物量比无显著差异.弱光环境促进杉木种子萌发,不利于杉木幼苗存活和生长.在弱光环境下,杉木幼苗通过增大茎生物量来提高对弱光环境的耐受力.