Cotyledon venation patterns in the Leguminosae: Gaesalpinioideae

Cotyledon venation patterns are described for 93 species representing all tribes of the Caesalpinioideae. Patterns are grouped into a series of levels of complexity according to the number of primary veins, nine, seven, six, five, three or one, but five- and three-vein patterns predominate. The number of petiolar vascular strands varies from one to eight but most species have two or four strands. It is proposed that all the patterns have been derived from one in which four strands and a protoxylem trace in the petiole branch and anastomose to form seven primary veins in the lamina. Venation patterns show correlations with cotyledon anatomy, size and shape and with taxonomic grouping at tribal, infratribal and generic levels. Each tribe is characterized by a particular set of related patterns, pattern frequencies and evolutionary trends.     

黄精种子萌发过程发育解剖学研究

采用石蜡切片技术对成熟黄精种子形态及萌发过程中的形态学变化及解剖结构特征进行了研究,以阐明黄精种子繁殖的生物学机制。结果显示:(1)成熟的黄精种子由外而内依次为种皮、胚乳和胚等3部分组成。其中种皮由一层木质化的细胞组成;胚乳占据种子的大部分结构,胚乳细胞含有大量淀粉,细胞壁增厚;胚处于棒型胚阶段。(2)黄精种子在萌发过程中棒型胚靠近种脐端分化为吸器、子叶联结和子叶鞘,靠近种孔的部位分化出胚根、胚轴和胚芽。(3)黄精种子萌发首先由子叶联结伸长将胚芽和胚根原基推出种孔,紧接着下胚轴膨大形成初生小根茎,吸器留在种子中分解吸收胚乳中的营养物质。(4)通过子叶联结连通吸器和初生小根茎,将胚乳中的营养物质由吸器-子叶联结这个通路转移到初生小根茎中,为初生根茎上胚芽和胚根的进一步分化提供物质保障。(5)黄精种子自然条件下萌发率较低,而且当年不出土。研究表明,黄精种子的繁殖生物学特性是其生态适应的一种重要机制。     

Cotyledon areoles in Bobgunnia (Fabaceae: Swartzieae)

Cotyledon areoles, a seed character unique to papilionoid legumes, are reported for the first time in the legume tribe Swartzieae, an anomalous group sometimes considered to be intermediate between subfamilies Caesalpinioideae and Papilionoideae. Bobgunnia madagascariensis has a linear, slightly branched cotyledon areole on the abaxial surface of each cotyledon. Previous reports of other papilionoid seed features in this species are confirmed and extended. These highly distinctive papilionoid seed structures, and recent reports from floral ontogeny and molecular systematics which show Bobgunnia to be closely related to Swartzia and several other genera of Swartzieae, add further evidence to assertions that Swartzia seed structures are derived by simplification from a papilionoid seed, and not by retention of less specialized caesalpinioid seed features. The function and evolutionary advantage of cotyledon areoles are unknown and need investigation. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 287–291.     

Cotyledon venation patterns in the Leguminosae: Mimosoideae

Cotyledon venation patterns are described for 131 species representing the four main tribes of the Mimosoideae. The range of variation in venation pattern falls within that described previously for the Caesalpinioideae and is consistent with the proposal that all patterns in the Leguminosae have been derived from a prototype with four vascular strands and a protoxylem trace in the petiole and seven primary veins in the lamina. Each tribe is characterized by a particular set of patterns, pattern frequencies and evolutionary trends. In the Mimoseae, correlations between venation pattern and cotyledon size and anatomy match those found in the Caesalpinioideae, but different correlations unique to these tribes occur in the Acacieae and Ingeae.     

Seed structure and the taxonomy of the Cruciferae

The external morphology and anatomy of the seeds of some 90 genera and some 200 species of the Cruciferae have been investigated. Emphasis has been laid on the relationships of seed structure to taxonomy. As regards seed anatomy, the testa is probably of greatest value in determining taxonomic relationships. Seed structure supports the Cruciferae as a natural taxon. In the present investigation, few genera and species show seed characters which are either exclusive or very distinct. Of the remainder, most genera are distinguishable from each other on a permutation of their macroscopic and microscopic seed characters. However it is difficult or impossible to distinguish between some genera on seed characters. The present investigation provides no support for tribal divisions within the Cruciferae.     

Endopolyploidy, Cell Volume and Nuclear Volume Interrelationships in Cotyledons of the Leguminosae

Cotyledon nuclei of most legume species did not stain adequatelywith the Feulgen reaction and so an indirect method based onthe relationship between DNA content, nuclear volume and cellvolume was used for the detection of endopolyploidy. Cotyledonendopolyploidy was not found in the Caesalpinioideae and wasinfrequent in the Mimosoideae. In the Papilionoideae it wasfound in taxa at several levels. It probably occurs throughoutthe tribes Vicieae and Cicereae and in most genera of the Phaseoleae;elsewhere it is confined to particular genera within tribes,such as Arachis in the Aeschynomeneae and Lupinus in the Genisteae.Endopolyploidy shows correlation with cotyledon anatomy aridfunction in that it is confined to cotyledons whose main oronly function is storage, and has so far not been found in foliarcotyledons. Endopolyploidy, cell volume, nuclear volume, cotyledon, Leguminosae     

Evolutionary and ecological correlates of early seedling morphology in East African trees and shrubs

Seed size and cotyledon morphology are two key juvenile traits that have evolved in response to changes in plant species life-history strategies and habitat associations. Correlations of these traits with each other and with other juvenile traits were examined for 70 species of trees and shrubs in Kibale National Park, Uganda. Although species with photosynthetic cotyledons were more abundant than in other tropical floras, both univariate and multivariate analyses supported trait associations expected from the literature. Trait values varied continuously across species, yet mean trait values differed significantly among habitat association types. Species with large seeds, large seedlings, thick storage cotyledons, slow germination, large-stature adults, and dispersal by large animals were common in forest and gap habitats. An opposite suite of traits was common in open habitats (grassland and edge). Analyses incorporating phylogeny (independent contrasts and omnibus tests) confirmed that these suites of traits showed correlated evolution. Cotyledon functional morphology yielded a strong phylogenetic signal, while seed mass was labile. Nevertheless, contingent change tests found that evolutionary change from photosynthetic to reserve cotyledons was more likely when disperser and perhaps seed size of ancestral species were already large, suggesting a strong interdependency among these traits.     

Cotyledon damage at the seedling stage affects growth and flowering potential in mature plants

Seedling herbivory is an important selective filter influencing patterns of plant community composition. Nevertheless, while many of the mechanisms governing seedling selection by herbivores are well established, the effects of tissue loss at the seedling stage on subsequent plant development are poorly understood. Here we examined how the removal of 50 or approximately 100% of cotyledon area from 7-d-old chalk grassland seedlings affected subsequent plant growth and flowering over a 100-d period. Cotyledon damage had a significant effect on growth during the establishment phase for six of the nine species. For two species, significant effects on plant growth were manifest in 100-d-old plants. Of the five species that flowered, three developed fewer inflorescences or flowered later as a consequence of cotyledon damage suffered as a seedling. Our results show that, in addition to the direct effect of herbivory on seedling mortality, more subtle sublethal effects may also influence plant establishment. Reduced growth as a result of cotyledon damage may have implications for plant competition during the establishment phase, and on subsequent reproductive success at maturity.     

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.     

Seedling diversity in Hydatellaceae: implications for the evolution of angiosperm cotyledons

BACKGROUND AND AIMS: Cotyledon number has long been a primary morphological feature distinguishing monocots from other angiosperms. Recent placement of Hydatellaceae near the early-divergent angiosperm order Nymphaeales, rather than in the monocot order Poales, has prompted reassessment of seedling morphology in this poorly known family. METHODS: Seedlings of six species representing all eco-geographical groups of Hydatellaceae are described using light and scanning electron microscopy. KEY RESULTS: Two seedling types were discovered. Material examined of Trithuria submersa, T. bibracteata, T. austinensis and T. filamentosa possess a transparent bilobed sheathing structure that surrounds the main axis below the first foliage leaf. The seed coat is attached to the sheathing structure. Seedlings of Trithuria lanterna and T. konkanensis lack a sheathing structure, and the seed coat is attached to a short, narrow lateral outgrowth on the main axis of the seedling. CONCLUSIONS: The sheathing structure that is present in seedlings of some Hydatellaceae could be homologized with the two united cotyledons of water lilies. It also resembles the single cotyledon of some monocots, and hence demonstrates a possible pathway of the origin of a monocot-like embryo, though no homology is implied. The sheathing structure is reduced in Trithuria lanterna and T. konkanensis, and the short, narrow outgrowth of its seedling could represent a single cotyledon. This synapomorphy suggests that the only Indian species of Hydatellaceae, T. konkanensis, is closer to the northern Australian T. lanterna than to the south-western Australian T. bibracteata.     

Parturition following transfer of embryos produced in two media under two oxygen concentrations

In vitro-produced blastocysts were transferred singly to 99 Angus crossbred recipients. Culture treatments were a 2 x 2 factorial of medium (KSOM or SOF) and oxygen concentration (5 or 20%). At parturition, birth weight and frame measurement before colostrum intake were recorded. Fetal membranes were collected; distribution and diameter of cotyledons was recorded. Cotyledon surface area was calculated. Culture with 5% O2 tended to yield smaller birth weights than culture with 20% O2 (39.7+/-1.3 kg versus 43.1+/-1.4 kg; P < 0.1); this effect was pronounced in KSOM, suggesting a medium by oxygen interaction (P < 0.1). When expressed on a body weight basis, calves born following culture with 20% O2 had consistently smaller skeletal measurements than those from culture with 5% O2. Culture with 20% O2 significantly increased individual cotyledon areas both overall and in the fetal horn and cotyledon surface area in the fetal horn. Overall, individual cotyledons were 32% larger when culture involved 20% O2 versus 5% O2; in the fetal horn the increase was 49%. Cotyledon surface area was greater for 20% compared to 5% O2 culture, though a medium by oxygen interaction was also significant (P < 0.05). Cotyledon surface area in the nonfetal horn was greater for KSOM fetal membranes than those from culture in SOF. There was a significant medium by oxygen interaction for total cotyledon number. These data demonstrate culture system-specific effects on calf and fetal membrane traits.     

Colic modifications in iguanine lizards

All lizard species of the subfamily Iguaninae except Amblyrynchus cristatus possess from one to eleven transverse valves in the proximal colon. Valves are of two kinds: circular (sometimes with a sphincter valve) or semilunar. Circular valves (if present) always occur proximally to semilunar valves. Intraspecific variation in the number and type of valves is small, but increase with modal number of valves. No significant ontogenetic change in number of valves could be demonstrated. Colic valves in iguanine lizards apparently evolved as simple infoldings of the medical colic wall. Comparisons are made with colic modifications occurring in other lizard families. Herbivorous species of the Scincidae, Agamidae, and Iguanidae are the only lizards known to exhibit colic partitioning, suggesting that the evolution of these structures is intimately related to the evolution of herbivory in these lizards. The potential taxonomic and phylogenetic importance of lizard colon anatomy is discussed.     

Timing of cotyledon damage affects growth and flowering in mature plants

Although the effects of herbivory on plant fitness are strongly linked to age, we understand little about how the timing of herbivory at the seedling stage affects growth and reproduction for plants that survive attack. In this study, we subjected six north-western European, dicotyledonous grassland species (Leontodon autumnalis, Leontodon hispidus, Plantago lanceolata, Plantago major, Trifolium pratense and Trifolium repens) to cotyledon removal at 7, 14 and 21 d old. We monitored subsequent growth and flowering (number of inflorescences recorded, and time taken for first flowers to open) over a 107 d period. Cotyledon removal reduced growth during establishment (35 d) for all species, and a further three exhibited reduced growth at maturity. Four species developed fewer inflorescences, or had delayed flowering after cotyledon removal. Although early damage (7 d old) had the greatest long-term effect on plant performance, responses varied according to the age at which the damage occurred and the species involved. Our results illustrate how growth and flowering into the mature phase is affected by cotyledon damage during different stages of seedling ontogeny, and we highlight the ways in which ontogenetic variation in seedling tolerance of tissue loss might impact upon plant fitness in mature plant communities.     

A Comparative Study of the Role of the Cotyledon in Seedling Development

Seedling development was studied in four species in which thecotyledons had different capacities for expansion and CO²-flxation. The highest growth-rates were shown by the two species in whichthe cotyledons showed the most expansion and also the greatest¹⁴CO²-fixation capacity. Cotyledons of these species becamevery leaf-like and retained a high proportion of the radiocarbonfixation products during their growth in contrast to the hypogealrunner-bean cotyledon which had a high percentage export valuealthough total fixation was low. Seedlings with leaf-like cotyledons which had the dual roleof storage and subsequent provision of photosynthetic organshad delayed leaf development with early growth concentratedin cotyledon, hypocotyl, and root whereas species with cotyledonsless well adapted for photosynthesis produced leaves at an earlierstage. Thus, the pattern of early seedling development was closelyrelated to the degree of photosynthetic adaptation shown bythe cotyledons. The evidence suggests that in the species studied, cotyledonsadapted for the dual role appear to provide a more efficientsystem for early seedling development.     

A survey of seed and seedling characters in 1744 Australian dicotyledon species: cross-species trait correlations and correlated trait-shifts within evolutionary lineages

Seedling traits have been described across 1744 species of Australian plants. Six traits were coded as binary alternatives: (1) phanerocotyly vs cryptocotyiy; (2) first leaf scale‐like vs leaf‐like; (3) first leaf single vs paired; (4) cotyledons hairy vs. glabrous; (5) hypocotyl hairy vs glabrous, and (6) embryo green vs non‐green at maturity. Seed volume was calculated from measurements of seed dimensions. Three approaches were used in analysing the data: (1) the taxonomic distribution of binary character states was described; (2) the strength of bivariate character associations was quantified at species level (‘cross‐species' correlations and regressions); (3) the data were arrayed on a phylogenetic tree in order to analyse by ‘phylogenetic regression’ for correlated evolutionary shifts in trait pairs. All die traits appeared evolutionarily malleable. For example, while cryptocotyiy was the minority condition (22% of species), it occurred in many different taxa (124 genera, 40 families, 24 orders) with high levels of polymorphy (16%, 25% and 42% for genera, families and orders). Similarly, the less common attribute states for first leaf type, hypocotyl texture, cotyledon texture and embryo colour occurred in species from right across die dicotyledon phylogeny, as did independent evolutionary divergences in each of mese characters. These patterns indicate that debate over which conditions are primitive and which are advanced will not have any general answer, only an answer for a specific branch‐step in die phylogenetic tree. In nearly all cases, correlated‐divergence analyses showed the same patterns as cross‐species analyses. The strongest associations were between seed volume and cryptocotyiy, seed volume and scale‐like first leaf, and seed volume and presence of green embryo. In addition, cryptocotyiy and scale‐like first leaf, and cotyledon and hypocotyl type, were strongly associated in bom correlated‐divergence and cross‐species analyses. Interpretation of results was mosdy presented wim respect to seed size, a trait which we consider to be pivotal in a species' seedling establishment strategy. All possible pairwise combinations ofbinary seedling traits were found in our study species. Taken togedier, the various lines of evidence presented here suggest that the traits have assorted more or less independendy of each odier and provide no evidence of functional groups based an mese attributes. Thus, the several existing seedling typologies should be regarded as classifications of convenience rather than as reflecting fundamental types.     

种子重量与海拔的相关性分析

种子重量与海拔的关系是植物生态学中重要而尚存争议的问题。本文对中国123科853种2879份野生植物种子的重量在海拔上的变化特点以及二者之间的相互关系进行了深入研究,结果显示：种子重量总体上与海拔呈负相关,种子重量级别多样性和生活型多样性随海拔上升呈阶梯状下降。但7级种子中,只有100级种子的重量与海拔呈负相关;7种生活型中,只有灌木和匍匐草本的种子重量与海拔呈负相关;在种水平上,只有82%的物种种子重量与海拔存在相关性。本研究揭示了我国种子重量与海拔之间的复杂关系,以及遗传和环境对种子重量的影响,对植物育种、森林管理以及入侵植物的防治具有一定指导意义。     

Determinism of carbon and nitrogen reserve accumulation in legume seeds

In legume plants, the determination of individual seed weight is a complex phenomenon that depends on two main factors. The first one corresponds to the number of cotyledon cells, which determines the potential seed weight as the cotyledon cell number is related to seed growth rate during seed filling. Since cell divisions take place between flowering and the beginning of seed filling, any stress occurring before the beginning of seed filling can affect individual seed growth rate (C and N reserve accumulation in seeds), and thus individual seed weights. The second factor concerns carbon and nitrogen supply to the growing seed to support reserve accumulation. Grain legume species produce protein-rich seeds involving high requirement of nitrogen. Since seed growth rate as determined by cotyledon cell number is hardly affected by photoassimilate availability during the filling period, a reduction of photosynthetic activity caused by nitrogen remobilization in leaves (e.g., remobilization of essential proteins involved in photosynthesis) can lead to shorten the duration of the filling period, and by that can provoke a limitation of individual seed weights. Accordingly, any biotic or abiotic stress during seed filling causing a decrease in photosynthetic activity should lead to a reduction of the duration of seed filling.     

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.     

Light requirements for regeneration in tropical forest plants: Taxon-level and ecological attribute effects

Abstract Understanding co-adapted traits and strategies in organisms has been increasingly shown to require a broader view of structure, function, taxonomic affiliation (phylogeny) and their possible interactions. In order to understand mechanisms of recruitment dynamics in tropical forest, nine juvenile (fruit, seed and seedling) and seven adult ecological and morphological traits were examined in 89 woody plant species, from 75 genera, 35 families and 21 orders. The species were from tropical rainforest in northern Queensland and varied widely in light requirements and hence regeneration niches. The importance of taxonomic (phylogenetic) constraints was considered using various techniques. Data were collected mostly from herbarium specimens, augmented by observations in the field. The main source of variation in most traits was at the species level, although fruit size and light requirement varied most at the genus level, and flower size at the level of order. Covariation among attributes increased in strength with increasing taxonomic level. Cluster analyses of the species arising either from five or three groups reflected juvenile rather than adult attributes. However, these groups were not morphologically distinct. Sets of juvenile traits poorly predict adult functions (and vice-versa) at species, genus and family levels, but do so more strongly at the order level. Analyses of variance and multivariate analyses, with and without the effects of phylogeny, indicated that fruit size, seed size, hardness of seed coat, seedling cotyledon function, fruit maturation period and, to a lesser extent, seed number per fruit and wood density could predict light requirement, and hence regeneration niche. Flower size showed a strong phylogenetic effect, and thus is conserved along taxonomic lines. It is concluded that as different traits may vary at different taxonomic levels, the influence of phylogeny on interspecific variation needs to be routinely recognized by ecologists involved in comparative analyses of plant life-history functions.