Climate and growth form: the consequences for genome size in plants

The adaptive significance of nuclear DNA variation in angiosperms is still widely debated. The discussion mainly revolves round the causative factors influencing genome size and the adaptive consequences to an organism according to its growth form and environmental conditions. Nuclear DNA values are now known for 3874 angiosperm species (including 773 woody species) from over 219 families (out of a total of 500) and 181 species of woody gymnosperms, representing all the families. Therefore, comparisons have been made on not only angiosperms, taken as a whole, but also on the subsets of data based on taxonomic groups, growth forms, and environment. Nuclear DNA amounts in woody angiosperms are restricted to less than 23.54 % of the total range of herbaceous angiosperms; this range is further reduced to 6.8 % when woody and herbaceous species of temperate angiosperms are compared. Similarly, the tropical woody dicots are restricted to less than 50.5 % of the total range of tropical herbaceous dicots, while temperate woody dicots are restricted to less than 10.96 % of the total range of temperate herbaceous dicots. In the family Fabaceae woody species account for less than 14.1 % of herbaceous species. Therefore, in the total angiosperm sample and in subsets of data, woody growth form is characterized by a smaller genome size compared with the herbaceous growth form. Comparisons between angiosperm species growing in tropical and temperate regions show highly significant differences in DNA amount and genome size in the total angiosperm sample. However, when only herbaceous angiosperms were considered, significant differences were obtained in DNA amount, while genome size showed a non-significant difference. An atypical result was obtained in the case of woody angiosperms where mean DNA amount of tropical species was almost 25.04 % higher than that of temperate species, which is because of the inclusion of 85 species of woody monocots in the tropical sample. The difference becomes insignificant when genome size is compared. Comparison of tropical and temperate species among dicots and monocots and herbaceous monocots taken separately showed significant differences both in DNA amount and genome size. In herbaceous dicots, while DNA amount showed significant differences the genome size varies insignificantly. There was a non-significant difference among tropical and temperate woody dicots. In three families, i.e., Poaceae, Asteraceae, and Fabaceae the temperate species have significantly higher DNA amount and genome size than the tropical ones. Woody gymnosperms had significantly more DNA amount and genome size than woody angiosperms, woody eudicots, and woody monocots. Woody monocots also had significantly more DNA amount and genome size than woody eudicots. Lastly, there was no significant difference between deciduous and evergreen hardwoods. The significance of these results in relation to present knowledge on the evolution of genome size is discussed.     

Nuclear DNA C-values Complete Familial Representation in Gymnosperms

The gymnosperms are a monophyletic yet diverse group of woodytrees with approx. 730 extant species in 17 families. A recentsurvey showed that DNA C-values were available for approx. 16%of species, but for only 12 of the 17 families. This paper completesfamilial representation reporting first C-values for the fiveremaining families: Boweniaceae, Stangeriaceae, Welwitschiaceae,Cephalotaxaceae and Sciadopityaceae. C-values for nine Ephedraand two Gnetum species are also reported. C-values are now availablefor 152 (21%) species. Analysis confirms that gymnosperms arecharacterized by larger C-values than angiosperms (modal 1Cof gymnosperms = 15.8 pg compared with 0.6 pg in angiosperms)although the range (1C = 2.25–32.20 pg) is smaller thanthat in angiosperms (1C = 0.05–127.4 pg). Given completefamilial coverage for C-values and increasing consensus in gymnospermphylogeny, the phylogenetic component of C-value variation wasalso investigated by comparing the two datasets. This analysisrevealed that ancestral gymnosperms (represented by cycads and/orGinkgo; mean genome size = 14.71 pg) probably had larger genomes thanancestral angiosperms. Copyright 2001 Annals of Botany Company Gymnosperm DNA amounts, C-values, phylogeny, ancestral genome size, Cycadales, Ginkgo, Gnetales, conifers, Pinaceae     

Accessory costs of seed production and the evolution of angiosperms

Accessory costs of reproduction frequently equal or exceed direct investment in offspring, and can limit the evolution of small offspring sizes. Early angiosperms had minimum seed sizes, an order of magnitude smaller than their contemporaries. It has been proposed that changes to reproductive features at the base of the angiosperm clade reduced accessory costs thus removing the fitness disadvantage of small seeds. We measured accessory costs of reproduction in 25 extant gymnosperms and angiosperms, to test whether angiosperms can produce small seeds more economically than gymnosperms. Total accessory costs scaled isometrically to seed mass for angiosperms but less than isometrically for gymnosperms, so that smaller seeds were proportionally more expensive for gymnosperms to produce. In particular, costs of abortions and packaging structures were significantly higher in gymnosperms. Also, the relationship between seed:ovule ratio and seed size was negative in angiosperms but positive in gymnosperms. We argue that the carpel was a key evolutionary innovation reducing accessory costs in angiosperms by allowing sporophytic control of pre- and postzygotic mate selection and timing of resource allocation. The resulting reduction in costs of aborting unfertilized ovules or genetically inferior embryos would have lowered total reproductive costs enabling early angiosperms to evolve small seed sizes and short generation times.     

Variation in the psbC gene region of gymnosperms and angiosperms as detected by a single restriction site polymorphism

 This paper reports on a PCR-RFLP analysis in a chloroplast DNA region consisting of coding and intergenic spacer sequences of trnS and the adjacent psbC gene. This region was PCR-amplified in 62 woody plant species, predominantly tree species, that represent a broad systematic range in both gymnosperms and dicotyledonous angiosperms. The amplification products were digested by the restriction endonuclease HaeIII (GG↓CC). Fourteen different restriction patterns occurred, 5 of which characterised representatives of the gymnosperms, and 9 angiosperm representatives. A single restriction site polymorphism revealed most of the species to share restriction patterns. Groups formed which showed relationships to plant systematic units. This phenomenon is discussed with regard to the psbC gene and the GGCC motif for tracing species’ relationships on a high taxonomic level of gymnosperms and angiosperms. Received: 5 June 1996/Accepted: 20 September 1996     

Sexual systems in gymnosperms: A review

The aim of this study was to update figures for the presence of dioecy among the gymnosperms and investigate its correlation with climate, growth form, pollination and seed dispersal syndromes, and risk of extinction. Dioecy was found in almost 65% of contemporary gymnosperm species, a higher percentage than previous estimates. It dominates in 8 of the 12 families. As in angiosperms, dioecious gymnosperms are particularly common in climbers and are more commonly found in tropical climates. Analysis of the degree of threat using IUCN red list categories showed that the proportion of threatened species is higher in dioecious than in monoecious species only in temperate climate. The high sensitivity of dioecious species to environmental changes associated with human activity in temperate climate may explain this phenomenon. The monophyly of extant gymnosperms and the relatively small number of species (about 1000) create the possibility of treating them as a model group in investigating the evolution of sexual systems.     

晋东南山地种子植物区系研究

晋东南山地有野生种子植物974种,隶属于456属125科,其中裸子植物7科11属15种,被子植物118科445属959种(其中双子叶植物104科396属833种,单子叶植物14科,49属126种)。种子植物属的区系成分复杂多样,以温带成分占绝对优势(温带属250属,占总属数的60．98％)。种子植物种的区系成分中,中国特有种占绝对优势,共有383种,占总种数的39．32％,它们是构成晋东南山地森林植被和灌丛植被的建群种和优势种的主要成分。     

Correlated evolution of genome size and seed mass

Previous investigators have identified strong positive relationships between genome size and seed mass within species, and across species from the same genus and family. Here, we make the first broad-scale quantification of this relationship, using data for 1222 species, from 139 families and 48 orders. We analyzed the relationship between genome size and seed mass using a statistical framework that included four different tests. A quadratic relationship between genome size and seed mass appeared to be driven by the large genome/seed mass gymnosperms and the many small genome size/large seed mass angiosperms. Very small seeds were never associated with very large genomes, possibly indicating a developmental constraint. Independent contrast results showed that divergences in genome size were positively correlated with divergences in seed mass. Divergences in seed mass have been more closely correlated with divergences in genome size than with divergences in other morphological and ecological variables. Plant growth form is the only variable examined thus far that explains a greater proportion of variation in seed mass than does genome size.     

山西云顶山自然保护区野生种子植物区系研究

山西云顶山自然保护区位于吕梁山脉中段,有野生种子植物926种,隶属于391属、85科,其中裸子植物2科、4属、6种,被子植物83科、387属、920种(其中双子叶植物72科、311属、756种;单子叶植物11科、76属、164种).种子植物属的区系成分复杂多样,以温带成分占绝对优势,温带属共275属,占总属数的82.08％.种子植物种的区系成分中,温带分布占绝对优势,达566种,占总种数的62.61％,中国特有分布次之,有306种,它们是构成云顶山自然保护区森林植被和灌丛植被的建群种和优势种的主要成分.经与有关山地植物区系和植被组成比较后得出,云顶山自然保护区与庞泉沟自然保护区具有较高的同质性和密切的联系,与五台山次之,而与中条山具有明显的差异.云顶山自然保护区有国家二级保护植物野大豆,还有山西省重点保护植物山西乌头、红景天和党参3种.     

江西野生种子植物区系多样性及其基本特征

江西位于我国中亚热带东部,地史悠久,自然环境条件优越,植物种类相当丰富.初步统计江西共有野生种子植物4057种,隶属于193科1082属,其中裸子植物7科18属29种,被子植物186科1064属4028种.科属区系研究表明:(1)区系组成丰富,优势科、属明显;(2)地理成分复杂,区系联系广泛,以热带性成分为主,温带性成分占有较大的比例;(3)区系起源古老,孑遗植物众多;(4)珍稀植物和特有植物丰富.     

Relative rates of synonymous substitutions in the mitochondrial, chloroplast and nuclear genomes of seed plants

Previous studies have estimated that, in angiosperms, the synonymous substitution rate of chloroplast genes is three times higher than that of mitochondrial genes and that of nuclear genes is twelve times higher than that of mitochondrial genes. Here we used 12 genes in 27 seed plant species to investigate whether these relative rates of substitutions are common to diverse seed plant groups. We find that the overall relative rate of synonymous substitutions of mitochondrial, chloroplast and nuclear genes of all seed plants is 1:3:10, that these ratios are 1:2:4 in gymnosperms but 1:3:16 in angiosperms and that they go up to 1:3:20 in basal angiosperms. Our results show that the mitochondrial, chloroplast and nuclear genomes of seed plant groups have different synonymous substitutions rates, that these rates are different in different seed plant groups and that gymnosperms have smaller ratios than angiosperms.     

山西万家寨引黄工程沿线种子植物区系分析

山西万家寨引黄工程从万家寨水利枢纽至太原市呼延水厂, 全长285.03 km 。引黄工程沿线有种子植物1 040 种, 隶属于422 属100 科, 其中裸子植物3 科5 属9 种;被子植物97 科417属1 031 种(双子叶植物79 科321 属793 种, 单子叶植物18 科96 属238 种)。引黄工程沿线种子植物属的区系成分类型多样, 具有明显的温带色彩。突出特征是温带成分占优势, 达246 属, 占总属数的58.29%(其中北温带成分155 属, 占总属数的36.73%), 在植物区系的组成中具有重要作用。在引黄工程沿线种子植物种的区系成分中, 中国特有种占绝对优势, 达311 种, 占总种数的29.90%, 它们是构成引黄工程沿线优势植被类型的建群种和优势种的主要成分, 如青、华北落叶松、油松、小叶杨、青杨、垂柳、旱榆和黄刺玫等。     

Pines as Model Gymnosperms To Study Evolution, Wood Formation, and Perennial Growth

Pines provide a model system for the gymnosperms, an old and successful group of vascular plants that last shared a common ancestor with the angiosperms about 285 million years ago. Gymnosperms are distinct from angiosperms in their reproduction, development, metabolism, adaptations, and evolution. Pines cover vast areas of the globe, are one of the most important genera of forest trees, dominate the ecology of many temperate and subtropical forest ecosystems, and provide a major fraction of the world's wood. Here, we summarize many features of pine that make it a useful model for gymnosperms and woody plants. We also describe the influence of its reproductive system on methods for genetic analysis and the prospects for genomic studies and genetic engineering. Pines are limited as model systems by their long generation times, large size, large genomes, and the long time from fertilization to seed set.     

屏山县老君山自然保护区种子植物区系的研究

通过野外调查和查阅资料,四川省屏山县老君山自然保护区有种子植物121科528属1 267种,其中裸子植物4科6属9种;被子植物117科522属1 258种。有39科种子植物属于泛热带分布类型,占总科数的44.83%;114属为北温带分布类型,占总属数的23.55%。保护区处于区系地域交汇处,与云南高原和滇黔桂地区属热带与亚热带的过渡性质的特征较不相符,而接近于横断山脉和华中、华东地区,区系表现为亚热带性质。     

Ecological and evolutionary determinants of a key plant functional trait: wood density and its community-wide variation across latitude and elevation

Wood density is an important trait in trees indicative of their life history and mechanical and physiological strategies. The following examines the evolutionary ecology of wood density using a large database for seed plants. In particular, we focused on the geographic and phylogenetic variation in wood density for both gymnosperms and angiosperms. A phylogenetic supertree was constructed for over 4600 taxa, allowing for comprehensive analyses of divergences across the seed plant phylogeny. Community-wide means and variances for wood densities were quantified for 171 standardized forest communities. Wood density was generally highly conserved across the entire seed plant phylogeny, yet large divergences were found within the rosid clade. Geographic and community variation in wood density, however, was significantly lower in temperate and high elevation communities, dominated by gymnosperms, than in tropical lowland communities, dominated by angiosperms, suggesting an increase in trait and, to some extent, clade filtering with latitude and elevation. Together, our results support the notion that both biotic and abiotic forces have been important in the evolution of wood density as well as in controlling the observed trait mean and variance across geographic gradients.     

Relationships between nuclear DNA content and seed and leaf size in soybean

 A correlation between genome size and agronomically important traits has been observed in many plant species. The goal of the present research was to determine the relationship between genome size, seed size, and leaf width and length in soybean [Glycine max (L.) Merr.] Twelve soybean strains, representing three distinct seed size groups, were analyzed. Flow cytometry was used to estimate their 2C nuclear DNA contents. Data on seed size and leaf size of the 12 strains were obtained from 1994 and 1995 field experiments. Variation of 2C nuclear DNA among the 12 soybean strains was 4.6%, ranging from 2.37 pg for a small-seed strain to 2.48 pg for a large-seed strain. Strain seed size was positively associated with leaf width (r=0.92) and leaf length (r=0.93). Genome size was highly correlated with seed size (r=0.97), leaf width (r=0.90) , and leaf length (r=0.93). The results of our study indicate that there is a significant correlation between genome size and leaf and seed size in soybean. It is possible that selection for greater seed size either leads to, or results from, greater genome size. If so, this relationship might be worth exploring at a more fundamental level. Received: 5 April 1997 / Accepted: 9 January 1998     

Genome size evolution in relation to leaf strategy and metabolic rates revisited

BACKGROUND AND AIMS: It has been proposed that having too much DNA may carry physiological consequences for plants. The strong correlation between DNA content, cell size and cell division rate could lead to predictable morphological variation in plants, including a negative relationship with leaf mass per unit area (LMA). In addition, the possible increased demand for resources in species with high DNA content may have downstream effects on maximal metabolic efficiency, including decreased metabolic rates. METHODS: Tests were made for genome size-dependent variation in LMA and metabolic rates (mass-based photosynthetic rate and dark respiration rate) using our own measurements and data from a plant functional trait database (Glopnet). These associations were tested using two metrics of genome size: bulk DNA amount (2C DNA) and monoploid genome size (1Cx DNA). The data were analysed using an evolutionary framework that included a regression analysis and independent contrasts using a phylogenetic tree with estimates of molecular diversification times. A contribution index for the LMA data set was also calculated to determine which divergences have the greatest influence on the relationship between genome size and LMA. KEY RESULTS AND CONCLUSIONS: A significant negative association was found between bulk DNA amount and LMA in angiosperms. This was primarily a result of influential divergences that may represent early shifts in growth form. However, divergences in bulk DNA amount were positively associated with divergences in LMA, suggesting that the relationship may be indirect and mediated through other traits directly related to genome size. There was a significant negative association between genome size and metabolic rates that was driven by a basal divergence between angiosperms and gymnosperms; no significant independent contrast results were found. Therefore, it is concluded that genome size-dependent constraints acting on metabolic efficiency may not exist within seed plants.     

The evolution of angiosperm seed proteins: A methionine-rich legumin subfamily present in lower angiosperm Clades

Analysis of legumin-encoding cDNAs fromDioscorea caucasica Lipsky (Dioscoreaceae) and fromAsarum europaeum L. (Aristolochiaceae) shows that there is an especially methionine-rich legumin subfamily present in the lower angiosperm clades including the Monocotyledoneae. It is characterized by a methionine content of 3–4 mol% which is roughly triple the methionine proportion of most other legumins. These “MetR” legumins, if present, still have to be detected in the higher angiosperms including the important seed crops. Evolutionary analysis suggests that the MetR legumins are the result of a gene duplication allowing the differentiation of legumin genes according to their sulfur content. The duplication event must have taken place before the split into mono- and dicotyledonous plants but probably after the separation of angiosperms and gymnosperms. Correspondence to: H. Fischer     

The tortoise and the hare: ecology of angiosperm dominance and gymnosperm persistence

Gymnosperms, and conifers in particular, are sometimes very productive trees yet angiosperms dominate most temperate and tropical vegetation. Current explanations for angiosperm success emphasize the advantages of insect pollination and seed dispersal by animals for the colonization of isolated habitats. Differences between gymnosperm and angiosperm reproductive and vegetative growth rates have been largely ignored. Gymnosperms are all woody, perennial and usually have long reproductive cycles. Their leaves are not as fully vascularized as those of angiosperms and are more stereotyped in shape and size. Gymnosperm tracheids are generally more resistant to solute flow than angiosperm vessels. A consequence of the less efficient transport system is that maximum growth rates of gymnosperms are lower than maximum growth rates of angiosperms in well lit, well watered habitats. Gymnosperm seedlings may be particularly uncompetitive since their growth depends on a single cohort of relatively inefficient leaves. Later, some gymnosperms attain a higher productivity than co-occurring angiosperm trees by accumulating several cohorts of leaves with a higher total leaf area. These functional constraints on gymnosperm growth rates suggest that gymnosperms will be restricted to areas where growth of angiosperm competitors is reduced, for example, by cold or nutrient shortages. Biogeographic evidence supports this prediction since conifers are largely confined to high latitudes and elevations or nutrient-poor soils. Experimental studies show that competition in the regeneration niche (between conifer seedlings and angiosperm herbs and shrubs) is common and significantly affects conifer growth and survival, Fast-growing angiosperms, especially herbs and shrubs, may also change the frequency of disturbance regimes thereby excluding slower-growing gymnosperms. Shade-tolerant and early successional conifers share similar characteristics of slow initial growth and low plasticity to a change in resources. Shade-tolerant gymnosperms would be expected to occur only where forest openings are small or otherwise unsuitable for rapid filling by fast-growing angiosperm trees, lianas or shrubs. The limited evidence available suggests that shade-tolerant conifers are confined to forests with small gap sizes where large disturbances are very rare. The regeneration hypothesis for gymnosperm exclusion by angiosperms is consistent with several aspects of the fossil record such as the early disappearance of gymnosperms from early successional environments where competition with angiosperms would have been most severe. However there are unresolved difficulties in interpreting process from paleoecological pattern which prevent the testing of alternative hypotheses.     

美姑大风顶自然保护区种子植物区系分析

四川美姑大风顶国家级自然保护区有种子植物 1 1 8科 ,3 85属 ,860种 ,其中裸子植物 5科 ,1 6属 ,1 9种 ;被子植物 1 1 3科 ,3 69属 ,841种。种子植物属、种的区系成分共有 1 3个类型 ;在所有成分中 ,以温带区系成分为主 ,达 1 81属 ,占总属数的 5 1 .5 8%。中国特有分布属共 1 1属 ,占总属数的 3 .1 3 % ;中国特有分布种共41 0种 ,占总种数的 48.41 %。保护区共有珍稀濒危植物 1 8种 ,其中国家一级保护植物 4种 ,国家二级保护植物 6种 ,国家三级保护植物 8种。     

横县野生种子植物区系及与附近地区的比较研究

为了解广西横县野生种子植物区系特征,对横县野生种子植物进行了调查分析。结果表明,广西横县有野生种子植物1 269种,隶属于163科658属,以被子植物占优势,而裸子植物则贫乏。在科属种组成上,以大科、区域单型属和少型属为主;生活型组成反映亚热带常绿阔叶林群落特征,藤本植物种类丰富。种子植物地理成分均以热带成分为主,兼备亚热带和温带成分。与邻近地区相比较,横县种子植物区系与广东紫金县相似性较高,区系成分组成则与广东新会区、紫金县相近。因此,横县地区野生种子植物种类丰富,热带性质明显,呈热带至亚热带过渡性质,起源古老,具有一定特有现象,与广东地区联系密切。