The cellular and molecular biology of conifer embryogenesis

Gymnosperms and angiosperms are thought to have evolved from a common ancestor c. 300 million yr ago. The manner in which gymnosperms and angiosperms form seeds has diverged and, although broad similarities are evident, the anatomy and cell and molecular biology of embryogenesis in gymnosperms, such as the coniferous trees pine, spruce and fir, differ significantly from those in the most widely studied model angiosperm Arabidopsis thaliana. Molecular analysis of signaling pathways and processes such as programmed cell death and embryo maturation indicates that many developmental pathways are conserved between angiosperms and gymnosperms. Recent genomics research reveals that almost 30% of mRNAs found in developing pine embryos are absent from other conifer expressed sequence tag (EST) collections. These data show that the conifer embryo differs markedly from other gymnosperm tissues studied to date in terms of the range of genes transcribed. Approximately 72% of conifer embryo-expressed genes are found in the Arabidopsis proteome and conifer embryos contain mRNAs of very similar sequence to key genes that regulate seed development in Arabidopsis. However, 1388 loblolly pine (Pinus taeda) embryo ESTs (11.4% of the collection) are novel and, to date, have been found in no other plant. The data imply that, in gymnosperm embryogenesis, differences in structure and development are achieved by subtle molecular interactions, control of spatial and temporal gene expression and the regulating agency of a few unique proteins.     

Cenozoic extinctions account for the low diversity of extant gymnosperms compared with angiosperms

We test the widely held notion that living gymnosperms are 'ancient' and 'living fossils' by comparing them with their sister group, the angiosperms. This perception derives partly from the lack of gross morphological differences between some Mesozoic gymnosperm fossils and their living relatives (e.g. Ginkgo, cycads and dawn redwood), suggesting that the rate of evolution of gymnosperms has been slow. We estimated the ages and diversification rates of gymnosperm lineages using Bayesian relaxed molecular clock dating calibrated with 21 fossils, based on the phylogenetic analysis of alignments of matK chloroplast DNA (cpDNA) and 26S nuclear ribosomal DNA (nrDNA) sequences, and compared these with published estimates for angiosperms. Gymnosperm crown groups of Cenozoic age are significantly younger than their angiosperm counterparts (median age: 32 Ma vs 50 Ma) and have long unbranched stems, indicating major extinctions in the Cenozoic, in contrast with angiosperms. Surviving gymnosperm genera have diversified more slowly than angiosperms during the Neogene as a result of their higher extinction rate. Compared with angiosperms, living gymnosperm groups are not ancient. The fossil record also indicates that gymnosperms suffered major extinctions when climate changed in the Oligocene and Miocene. Extant gymnosperm groups occupy diverse habitats and some probably survived after making adaptive shifts.     

Evolution of Reproductive Organs in Land Plants

LEAFY gene is the positive regulator of the MADS-box genes in flower primordia. The number of MADS-box genes presumably increased by gene duplications before the divergence of ferns and seed plants. Most MADS-box genes in ferns are expressed similarly in both vegetative and reproductive organs, while in gymnosperms, some MADS-box genes are specifically expressed in reproductive organs. This suggests that (1) the increase in the number of MADS-box genes and (2) the subsequent recruitment of some MADS-box genes as homeotic selector genes were important for the evolution of complex reproductive organs. The phylogenetic tree including both angiosperm and gymnosperm MADS-box genes indicates the loss of the A-function genes in the gymnosperm lineage, which is presumably related to the absence of perianths in extant gymnosperms. Comparison of expression patterns of orthologous MADS-box genes in angiosperms, Gnetales, and conifers supports the sister relationship of Gnetales and conifers over that of Gnetales and angiosperms predicted by phylogenetic trees based on amino acid and nucleotide sequences. Received 30 July 1999/ Accepted in revised form 9 September 1999     

Leaf evolution in Southern Hemisphere conifers tracks the angiosperm ecological radiation

The angiosperm radiation has been linked to sharp declines in gymnosperm diversity and the virtual elimination of conifers from the tropics. The conifer family Podocarpaceae stands as an exception with highest species diversity in wet equatorial forests. It has been hypothesized that efficient light harvesting by the highly flattened leaves of several podocarp genera facilitates persistence with canopy-forming angiosperms, and the angiosperm ecological radiation may have preferentially favoured the diversification of these lineages. To test these ideas, we develop a molecular phylogeny for Podocarpaceae using Bayesian-relaxed clock methods incorporating fossil time constraints. We find several independent origins of flattened foliage types, and that these lineages have diversified predominantly through the Cenozoic and therefore among canopy-forming angiosperms. The onset of sustained foliage flattening podocarp diversification is coincident with a declining diversification rate of scale/needle-leaved lineages and also with ecological and climatic transformations linked to angiosperm foliar evolution. We demonstrate that climatic range evolution is contingent on the underlying state for leaf morphology. Taken together, our findings imply that as angiosperms came to dominate most terrestrial ecosystems, competitive interactions at the foliar level have profoundly shaped podocarp geography and as a consequence, rates of lineage diversification.     

Ontogenetic shifts in plant–plant interactions in a rare cycad within angiosperm communities

Gymnosperms and angiosperms can co-occur within the same habitats but key plant traits are thought to give angiosperms an evolutionary competitive advantage in many ecological settings. We studied ontogenetic changes in competitive and facilitative interactions between a rare gymnosperm (Dioon sonorense, our target species) and different plant and abiotic neighbours (conspecific-cycads, heterospecific-angiosperms, or abiotic-rocks) from 2007 to 2010 in an arid environment of northwestern Mexico. We monitored survival and growth of seedlings, juveniles, and adults of the cycad Dioon sonorense to evaluate how cycad survival and relative height growth rate (RHGR) responded to intra- and interspecific competition, canopy openness, and nearest neighbour. We tested spatial associations among D. sonorense life stages and angiosperm species and measured ontogenetic shifts in cycad shade tolerance. Canopy openness decreased cycad survival while intraspecific competition decreased survival and RHGR during early ontogeny. Seedling survival was higher in association with rocks and heterospecific neighbours where intraspecific competition was lower. Shade tolerance decreased with cycad ontogeny reflecting the spatial association of advanced stages with more open canopies. Interspecific facilitation during early ontogeny of our target species may promote its persistence in spite of increasing interspecific competition in later stages. We provide empirical support to the long-standing assumption that marginal rocky habitats serve as refugia from angiosperm competition for slow-growing gymnosperms such as cycads. The lack of knowledge of plant–plant interactions in rare or endangered species may hinder developing efficient conservation strategies (e.g. managing for sustained canopy cover), especially under the ongoing land use and climatic changes.     

Leaf energy balance modelling as a tool to infer habitat preference in the early angiosperms

Despite more than a century of research, some key aspects of habitat preference and ecology of the earliest angiosperms remain poorly constrained. Proposed growth ecology has varied from opportunistic weedy species growing in full sun to slow-growing species limited to the shaded understorey of gymnosperm forests. Evidence suggests that the earliest angiosperms possessed low transpiration rates: gas exchange rates for extant basal angiosperms are low, as are the reconstructed gas exchange rates for the oldest known angiosperm leaf fossils. Leaves with low transpirational capacity are vulnerable to overheating in full sun, favouring the hypothesis that early angiosperms were limited to the shaded understorey. Here, modelled leaf temperatures are used to examine the thermal tolerance of some of the earliest angiosperms. Our results indicate that small leaf size could have mitigated the low transpirational cooling capacity of many early angiosperms, enabling many species to survive in full sun. We propose that during the earliest phases of the angiosperm leaf record, angiosperms may not have been limited to the understorey, and that some species were able to compete with ferns and gymnosperms in both shaded and sunny habitats, especially in the absence of competition from more rapidly growing and transpiring advanced lineages of angiosperms.     

Chilling rather than photoperiod controls budburst for gymnosperm species in subtropical China

低温而不是光周期调控中国亚热带裸子植物的出芽物候 摘要：被子植物春季物候的调控机制已经得到了广泛的研究。然而,裸子植物和被子植物在3亿年前就产生分化,裸子植物与被子植物的物候可能是受不同的因素所调控。亚热带植物物候的调节机制在很大程度上尚不明确,亚热带裸子植物物候是否由冷激需求和光照调控仍未得到验证。本研究在人工气候箱中设置了3个冷激处理和3 个光周期处理,并对切枝的出芽期进行了为期8周的研究。实验中我们测试了8种裸子植物：柳杉(Cryptomeria japonica)、杉木(Cunninghamia lanceolata)、柏树(Cupressus funebris)、银杏(Ginkgo biloba)、水杉(Metasequoia glyptostroboides)、马尾松(Pinus massoniana)、金钱 松(Pseudolarix amabilis)和罗汉松(Podocarpus macrophyllus),检验其出芽物候是否对光周期敏感或者是否具有较强的冷激需求,以及这两种因素哪个对促进出芽更为重要。研究结果表明,对于裸子植物,冷 激促进了出芽并提高了出芽率,而且裸子植物需要适度的低温天数来实现出芽。有趣的是,在同一森 林中裸子植物比被子植物对积温的需求更高。与德国温带裸子植物(194–600 d · °C)相比,亚热带裸子植 物(814–1150 d · °C)对积温的需求更高。光周期对裸子植物出芽的影响较小,说明冷激对裸子植物出芽的 促进作用大于光周期。这些结果表明,随着全球气候持续变暖,冬季气温的升高不仅会影响亚热带被子植物也会影响裸子植物的物候,从而可能导致春季出芽期的延迟。     

南昌市不同植物类群叶片氮磷浓度及其化学计量比

对南昌大学前湖校区89种主要植物叶片的N、P浓度及其化学计量比进行了研究,结果表明:乔灌、常绿、针叶、种子、裸子和单子叶植物类群的N浓度分别低于相对应的草本、落叶、阔叶、蕨类、被子和双子叶植物类群,而C3和C4植物差异不显著;乔灌、常绿和裸子植物类群的P浓度含量分别低于相对应的草本、落叶和被子植物类群,而针叶和阔叶、蕨类和种子、单子叶和双子叶、C3和C4植物类群间差异不显著;乔木、阔叶、被子和双子叶植物类群叶片N/P分别高于相对应的灌草、针叶、裸子和单子叶植物类群,而常绿和落叶、蕨类和种子、C3和C4植物类群之间差异不显著.可见,不同类型植物对N和P的吸收利用存在差异,且对不同养分供应采取不同的适应对策.结合研究区土壤养分现状,建议优先选择常绿、针叶、裸子和单子叶植物类群作为城市园林植物.     

The rise to dominance of the angiosperm kingdom: dispersal,habitat widening and evolution during the Late Cretaceous of Europe

Coiffard, C. & Gomez, B. 2009: The rise to dominance of the angiosperm kingdom: dispersal, habitat widening and evolution during the Late Cretaceous of Europe. Lethaia, Vol. 43, pp. 164–169. The earliest fossil records of angiosperms in Europe occur in the Barremian and consist of freshwater wetland plants. From the Barremian onwards, angiosperms show a stepwise widening of their ecological range with the result that they inhabited most environments by the Cenomanian. Nevertheless, most angiosperms had still restricted habitats, while a few angiosperm trees were confined to disturbed environments, such as channel margins. A Wagner’s Parsimony Method analysis performed on a fossil plant and locality database from the Turonian to the Campanian of Europe indicates continued decrease in richness of ferns and gymnosperms compared with angiosperms, turnover between conifer and palm trees in freshwater‐related swamps at about the Cenomanian/Turonian boundary, and spreading of angiosperm trees through the floodplains. The ecological range of angiosperm trees was increased, being recorded in channel margins from the Cenomanian and spreading over floodplains (e.g. Platanaceae) and swamps (e.g. Arecaceae) by the Campanian. These new ecological ranges and successions went with innovative architectures, such as dicot trees and palm trees. Most living core angiosperm families had their earliest representatives in the Late Cretaceous, which should be considered as the dawn of modern angiosperm forests. □Core angiosperms, Europe, Late Cretaceous, palms, Wagner’s Parsimony Method.     

Mesozoic plants and the problem of angiosperm ancestry

Krassilov, V.: Mesozoic plants and the problem of angiosperm ancestry.
Trends leading to the foliar and floral structures of angiosperms may be deduced by comparison with Mesozoic gymnosperms. The Debeya-Fontainea group of Cretaceous angiosperms closely resembles the Early Mesozoic Scoresbya group of pteridosperms with regard to leaf characters. The bivalved capsules of Jurassic Leptostrobus , with stigmatic bands, are regarded as the forerunners of certain types of angiosperm carpels. The angiospermous characters arose in several lineages of gymnosperms and were probably accumulated by non-sexual transfer of genetic material. The earliest angiosperm mega- and microfossils have been reported from the Middle and Upper Jurassic of the northern hemisphere. Most of these angiosperms were confined to chaparral-like communities dominated by shrubby conifers and cycadophytes. The rise of angiosperms was promoted by the climatic changes and the simultaneous rise of mammals.     

The evolution of gymnosperms redrawn by phytochrome genes: the Gnetatae appear at the base of the gymnosperms

Gymnosperms possess two to four phytochrome types which apparently are the result of successive gene duplications in the genomes of their common ancestors. Phytochromes are nuclear-encoded proteins whose genes, contrary to chloroplast, mitochondrion, and rRNA genes, have hitherto rarely been used to examine gymnosperm phylogenies. Since the individual phytochrome gene types implied phylogenies that were not completely congruent to one another, conflicting branching orders were sorted by the number of gene lineages present in a taxon. The Gnetatae (two gene types) branched at the base of all gymnosperms, a position supported by bootstrap sampling (distance and character state trees, maximum likelihood). The Gnetatae were followed by Ginkgo, Cycadatae, and Pinaceae (three gene types) and the remaining conifers (four gene types). Therefore, in phytochrome trees, the most ancient branch of the conifers (Pinatae) seems to be the Pinaceae. The next split appears to have separated Araucariaceae plus Podocarpaceae from the Taxaceae/Taxodiaceae/Cupressaceae group. Structural arrangements in the plastid genomes (Raubeson and Jansen 1992) corroborate the finding that there is no close connection between Pinaceae and Gnetatae as suggested by some publications. The analyses are based on 60 phytochrome genes (579 positions in an alignment of PCR fragments) from 28 species. According to rough divergence time estimates, the last common ancestor of gymnosperms and angiosperms is likely to have existed in the Carboniferous.     

Diversity and significance of late cretaceous permineralized plant remains from Hokkaido,Japan

Middle to Late Cretaceous permineralized plants hitherto described from Hokkaido, Japan are summarized. The fossil flora comprises fungi, ferns, gymnosperms and angiosperms. Many modern fern families have been recognized including Anemiaceae, Cyatheaceae, Dennstaedtiaceae, Gleicheniaceae Loxsomaceae, Lygodiaceae and Matoniaceae. Gymnosperms are most abundant in the flora. Some recently-found materials are tentatively introduced with brief comments emphasizing their morphological and taxonomical significance. A bisporangiate flower ofCycadeoidella japonica Ogura shows fine internal anatomy and provides evidence that the cycadeoidalean ovule was a cupulate, unitegmic structure. Vascular tracheids in the synangial wall support the evolution of cycadeoidalean synangia from Paleozoic seed-fern synangia. A new gymnosperm female fructification has a thick envelope comparable to an angiosperm carpel around a large seed. The angiosperms contain various morphologies that require further extensive study.     

Leaf economic traits from fossils support a weedy habit for early angiosperms

Many key aspects of early angiosperms are poorly known, including their ecophysiology and associated habitats. Evidence for fast-growing, weedy angiosperms comes from the Early Cretaceous Potomac Group, where angiosperm fossils, some of them putative herbs, are found in riparian depositional settings. However, inferences of growth rate from sedimentology and growth habit are somewhat indirect; also, the geographic extent of a weedy habit in early angiosperms is poorly constrained. Using a power law between petiole width and leaf mass, we estimated the leaf mass per area (LMA) of species from three Albian (110-105 Ma) fossil floras from North America (Winthrop Formation, Patapsco Formation of the Potomac Group, and the Aspen Shale). All LMAs for angiosperm species are low (<125 g/m(2); mean = 76 g/m(2)) but are high for gymnosperm species (>240 g/m(2); mean = 291 g/m(2)). On the basis of extant relationships between LMA and other leaf economic traits such as photosynthetic rate and leaf lifespan, we conclude that these Early Cretaceous landscapes were populated with weedy angiosperms with short-lived leaves (<12 mo). The unrivalled capacity for fast growth observed today in many angiosperms was in place by no later than the Albian and likely played an important role in their subsequent ecological success.     

Angiosperm origin and early stages of seed plant evolution deduced from rRNA sequence comparisons

Summary Complete or partial nucleotide sequences of five different rRNA species, coded by nuclear (18S, 5.8S, and 5S) or chloroplast genomes (5S, 4.5S) from a number of seed plants were determined. Based on the sequence data, the phylogenetic dendrograms were built by two methods, maximum parsimony and compatibility. The topologies of the trees for different rRNA species are not fully congruent, but they share some common features. It may be concluded that both gymnosperms and angiosperms are monophyletic groups. The data obtained suggest that the divergence of all the main groups of extant gymnosperms occurred after the branching off of the angiosperm lineage. As the time of divergence of at least some of these gymnosperm taxa is traceable back to the early Carboniferous, it may be concluded that the genealogical splitting of gymnosperm and angiosperm lineages occurred before this event, at least 360 million years ago, i.e., much earlier than the first angiosperm fossils were dated. Ancestral forms of angiosperms ought to be searched for among Progymnospermopsida. Genealogical relationships among gymnosperm taxa cannot be deduced unambiguously on the basis of rRNA data. The only inference may be that the taxon Gnetopsida is an artificial one, andGnetum andEphedra belong to quite different lineages of gymnosperms. As to the phylogenetic position of the two Angiospermae classes, extant monocotyledons seem to be a paraphyletic group located near the root of the angiosperm branch; it emerged at the earliest stages of angiosperm evolution. We may conclude that either monocotyledonous characters arose independently more than once in different groups of ancient Magnoliales or that monocotyledonous forms rather than dicotyledonous Magnoliales were the earliest angiosperms. Judging by the rRNA trees, Magnoliales are the most ancient group among dicotyledons. The most ancient lineage among monocotyledons leads to modern Liliaceae.     

Ecological and genetic factors linked to contrasting genome dynamics in seed plants

The large-scale replacement of gymnosperms by angiosperms in many ecological niches over time and the huge disparity in species numbers have led scientists to explore factors (e.g. polyploidy, developmental systems, floral evolution) that may have contributed to the astonishing rise of angiosperm diversity. Here, we explore genomic and ecological factors influencing seed plant genomes. This is timely given the recent surge in genomic data. We compare and contrast the genomic structure and evolution of angiosperms and gymnosperms and find that angiosperm genomes are more dynamic and diverse, particularly amongst the herbaceous species. Gymnosperms typically have reduced frequencies of a number of processes (e.g. polyploidy) that have shaped the genomes of other vascular plants and have alternative mechanisms to suppress genome dynamism (e.g. epigenetics and activity of transposable elements). Furthermore, the presence of several characters in angiosperms (e.g. herbaceous habit, short minimum generation time) has enabled them to exploit new niches and to be viable with small population sizes, where the power of genetic drift can outweigh that of selection. Together these processes have led to increased rates of genetic divergence and faster fixation times of variation in many angiosperms compared with gymnosperms.     

Light-dependent chloroplast development and expression of a light-harvesting chlorophyll a/b-binding protein gene in the gymnosperm Ginkgo biloba.

Unlike conifers, the gymnosperm Ginkgo biloba is dependent on light for chlorophyll (Chl) synthesis and initiation of chloroplast development. Dark-grown seedlings show complete etiolation, including no detectable Chl accumulation, no leaf expansion, and increased hypocotyl elongation. When dark-grown seedlings are placed in white light, Chl synthesis and leaf expansion are initiated, but unlike angiosperms, which initiate rapid photomorphogenesis, Ginkgo takes at least 1 week to change to a normal light-regulated pattern of growth. A cDNA clone (pLhcb*Gb1) encoding a Chl a/b-binding protein of light-harvesting complex II from Ginkgo mRNA has been used as a probe for the expression of this family of mRNAs. We have found that, in common with angiosperms but in marked contrast to pines, Lhcb mRNA is expressed in a highly light-dependent manner. In addition to being expressed in light-grown leaves, this sequence is also expressed in the green tissues of immature seeds. The Lhcb mRNA appears during greening in parallel with the onset of Chl synthesis. The complete sequence of pLhcb*Gb1 has been determined and the deduced amino acid sequence was found to be of type I based on comparison with signature sequences of angiosperm and gymnosperm sequences.     

Trait syndromes among North American trees are evolutionarily conserved and show adaptive value over broad geographic scales

Adaptive syndromes and their evolutionary constraints represent a powerful construct for understanding plant distributions. However, it is unclear how the species requirements to face multiple stressors promotes syndrome formation and to which abiotic stressors these syndromes show adaptive value over broad geographic scales. We combined local occurrence data from the U.S. Forest Inventory and Analysis (FIA) of 219 angiosperm and 85 gymnosperm species living across the conterminous US with phylogenies and trait data to identify tree syndromes, their evolutionary conservatism, and their adaptive value over broad scales. Factor analyses and evolutionary model selection revealed that trees possess functional trait syndromes that are strongly conserved. Major syndromes at the species level differed between angiosperms and gymnosperms. While the two main syndromes in angiosperms were related to cold and drought‐waterlogging tolerance, in gymnosperms a trade‐off between shade and drought tolerance was the main syndrome followed by a growth‐fire resistance syndrome. Additional RLQ and fourth‐corner approaches revealed that trait syndromes at the community level were broadly similar to those observed at the species level for angiosperms, although this was less clear for gymnosperms. This suggests that syndrome evolution has played an important role on angiosperm distributions, whereas additional ecological factors explain gymnosperm distributions. Importantly, syndromes show adaptive value, as they were geographically associated with several environmental variables showing structure from continental to local scales, being temperature the main abiotic stressor. Our results indicate that across the conterminous US tree species possess clear syndromes that are subjected to strong evolutionary constraints driving tree species and forest community distribution.     

Evolution of a unique anatomical precision in angiosperm leaf venation lifts constraints on vascular plant ecology

The main role of leaf venation is to supply water across the photosynthetic surface to keep stomata open and allow access to atmospheric CO₂ despite evaporative demand. The optimal uniform delivery of water occurs when the distance between veins equals the depth of vein placement within the leaf away from the evaporative surface. As presented here, only angiosperms maintain this anatomical optimum across all leaf thicknesses and different habitats, including sheltered environments where this optimization need not be required. Intriguingly, basal angiosperm lineages tend to be underinvested hydraulically; uniformly high optimization is derived independently in the magnoliids, monocots and core eudicots. Gymnosperms and ferns, including available fossils, are limited by their inability to produce high vein densities. The common association of ferns with shaded humid environments may, in part, be a direct evolutionary consequence of their inability to produce hydraulically optimized leaves. Some gymnosperms do approach optimal vein placement, but only by virtue of their ability to produce thick leaves most appropriate in environments requiring water conservation. Thus, this simple anatomical metric presents an important perspective on the evolution and phylogenetic distribution of plant ecologies and further evidence that the vegetative biology of flowering plants—not just their reproductive biology—is unique.