Why did heterospory evolve?

The primitive land plant life cycle featured the production of spores of unimodal size, a condition called homospory. The evolution of bimodal size distributions with small male spores and large female spores, known as heterospory, was an innovation that occurred repeatedly in the history of land plants. The importance of desiccation‐resistant spores for colonization of the land is well known, but the adaptive value of heterospory has never been well established. It was an addition to a sexual life cycle that already involved male and female gametes. Its role as a precursor to the evolution of seeds has received much attention, but this is an evolutionary consequence of heterospory that cannot explain the transition from homospory to heterospory (and the lack of evolutionary reversal from heterospory to homospory). Enforced outcrossing of gametophytes has often been mentioned in connection to heterospory, but we review the shortcomings of this argument as an explanation of the selective advantage of heterospory. Few alternative arguments concerning the selective forces favouring heterospory have been proposed, a paucity of attention that is surprising given the importance of this innovation in land plant evolution. In this review we highlight two ideas that may lead us to a better understanding of why heterospory evolved. First, models of optimal resource allocation – an approach that has been used for decades in evolutionary ecology to help understand parental investment and other life‐history patterns – suggest that an evolutionary increase in spore size could reach a threshold at which small spores yielding small, sperm‐producing gametophytes would return greater fitness per unit of resource investment than would large spores and bisexual gametophytes. With the advent of such microspores, megaspores would evolve under frequency‐dependent selection. This argument can account for the appearance of heterospory in the Devonian, when increasingly tall and complex vegetative communities presented competitive conditions that made large spore size advantageous. Second, heterospory is analogous in many ways to anisogamy. Indeed, heterospory is a kind of re‐invention of anisogamy within the context of a sporophyte‐dominant land plant life cycle. The evolution of anisogamy has been the subject of important theoretical and empirical investigation. Recent work in this area suggests that mate‐encounter dynamics set up selective forces that can drive the evolution of anisogamy. We suggest that similar dispersal and mating dynamics could have underlain spore size differentiation. The two approaches offer predictions that are consistent with currently available data but could be tested far more thoroughly. We hope to re‐establish attention on this neglected aspect of plant evolutionary biology and suggest some paths for empirical investigation.     

A quantitative comparison of foliage development among allopatric ferns,Dryopteris crassirhizoma,D. coreano-montana andD. filix-mas

A quantitative comparison was conducted on the foliage development during sporophyte development of three allopatric ferns in cool temperate and subalpine regions of Hokkaido and Tirol, European Alps. The foliage development ofDryopteris crassirhizoma, D. coreano-montana andD. filix-mas was quantitatively described by the leaf development (NV, number of veins); NV correlates the leaf-shape complexity from a circle (DI, L/2(3.14×S)^1/2). Nearly similar patterns were detected on frequency distribution of fertile leaves, fertility increase and number of leaves in threeDryopteris ferns which exhibit funnel-shaped foliage arrangements in mature sporophyte. No difference was found in number of leaves, maximum NV, fertility rate and leaf-shape parameters among three ferns. A positive difference was found only on changes in order of pinnae with maximum number of costa branches (NVP) and the DI of outline of pinnae betweenD. crassirhizoma andD. filix-mas. These allopatricDryopteris ferns seem to have a similar foliage structure, in spite of some sympatricDryopteris ferns capable of producing putative hybrids (D. austriaca andD. amurensis; D. tokyoensis andD. monticola) having different foliage structures in Hokkaido. Contribution No. 3346 from the Institute of Low Temperature Science, Hokkaido University.     

Molecular evolution of pteridophytes and their relationship to seed plants: Evidence from complete 18S rRNA gene sequences

Complete 18S ribosomal RNA sequence data from representatives of all extant pteridophyte lineages together with RNA sequences from different seed plants were used to infer a molecular phylogeny of vascular plants that included all major land plant lineages. The molecular data indicate that lycopsids are monophyletic and are the earliest diverging group within the vascular land plants, whereasPsilotum nudum is more closely related to the seed plants than to other pteridophyte lineages. The phylogenetic trees based on maximum likelihood, parsimony and distance analyses show substantial agreement with the evolutionary relationships of land plants as interpreted from the fossil record.     

Structure and function at the root apex - phylogenetic and ontogenetic perspectives on apical cells and quiescent centres

The summit of roots of various plant species may be occupied by a single, rapidly proliferating tetrahedral apical cell (as in ferns), or by a multicellular and slowly proliferating quiescent centre (as in angiosperms), or by intermediate types of cellular organizations. The present paper attempts to deduce the phylogeny of these various types of cellular patterning at the root apex.     

Restriction fragment length polymorphisms distinguish among accessions ofCeratopteris thalictroides andC. richardii (Parkeriaceae)

We have used cDNA clones as probes on Southern blots to detect restriction fragment length polymorphisms among sevenCeratopteris thalictroides accessions, threeC. richardii accessions, and one putative interspecific hybrid. We found that the stringency of post-hybridization washes was a critical parameter affecting the quality of our blots; even with homologous cDNA sequences low stringency conditions resulted in a smear of signal, but high stringency washes gave blots with distinct bands. Most probes showed hybridization with four or more genomic fragments. Similarities in the number and size of fragments between and within species indicated that (i)C. richardii shows limited polymorphism among accessions tested, (ii)C. thalictroides is highly polymorphic, and (iii) Hawaiian accessions ofC. thalictroides are divergent relative to their continental cohorts and among themselves. The putative interspecific hybrid did not group closely with either of these species.     

High diversity and low specificity of fungi associated with seedless epiphytic plants

Epiphytes, which grow on other plants for support, make up a large portion of Earth's plant diversity. Like other plants, their surfaces and interiors are colonized by diverse assemblages of fungi that can benefit their hosts by increasing tolerance for abiotic stressors and resistance to disease or harm them as pathogens. Fungal communities associated with epiphytic plants and the processes that structure these communities are poorly known. To address this, we sampled seven epiphytic seedless plant taxa in a Costa Rican rainforest and examined the effects of host identity and microhabitat on external and endophytic fungal communities. We found low host specificity for both external and endophytic fungi and weak differentiation between epiphytic and neighboring epilithic plant hosts. High turnover in fungi within and between hosts and habitats reveals that epiphytic plant-associated fungal communities are highly diverse and suggests that they are structured by stochastic processes.     

Development of secondary pine forests after pine wilt disease in western Japan

Abstract. The development of secondary Pinus densiflora (Japanese red pine) forests after pine wilt disease was studied through phytosociological analysis, estimation of forest structure before disease and size-structure, tree ring and stem analyses. Following the end of the disease, the growth of previously suppressed small oak trees was accelerated. This is quite different from the development of forests following fire, which starts with the establishment of pine seedlings. Pine wilt disease shifted the dominance of secondary forests from Pinus densiflora to Quercus serrata oak forest. In pine forests, disturbance by fire is important for forest maintenance. In contrast, disturbance by pine wilt disease leads to an acceleration of succession from pine forest to oak forest.     

Divergence of the phytochrome gene family predates angiosperm evolution and suggests thatSelaginella andEquisetum arose prior toPsilotum

Thirty-two partial phytochrome sequences from algae, mosses, ferns, gymnosperms, and angiosperms (11 of them newly released ones from our laboratory) were analyzed by distance and characterstate approaches (PHYLIP, TREECON, PAUP). In addition, 12 full-length sequences were analyzed. Despite low bootstrap values at individual internal nodes, the inferred trees (neighbor joining, Fitch, maximum parsimony) generally showed similar branching orders consistent with other molecular data. Lower plants formed two distinct groups. One basal group consisted of Selaginella, Equisetum, and mosses; the other consisted of a monophyletic cluster of frond-bearing pteridophytes. Psilotum was a member of the latter group and hence perhaps was not, as sometimes suggested, a close relative of the first vascular plants. The results further suggest that phytochrome gene duplication giving rise to a- and b- and later to c-types may have taken place within seedfern genomes. Distance matrices dated the separation of mono- and dicotyledons back to about 260 million years before the present (Myr b.p.) and the separation of Metasequoia and Picea to a fossil record-compatible value of 230 Myr B.P. The Ephedra sequence clustered with the c- or a-type and Metasequoia and Picea sequences clustered with the b-type lineage. The paleoherb Nymphaea branched off from the c-type lineage prior to the divergence of mono- and dicotyledons on the a- and b-type branches. Sequences of Piper (another paleoherb) created problems in that they branched off from different phytochrome lineages at nodes contradicting distance from the inferred trees' origin. Correspondence to: H.A.W. Schneider-Poetsch     

西双版纳热带雨林中两种生态型蕨类植物的光合特性比较研究

通过比较分布于西双版纳热带雨林林下生境中的附生鸟巢蕨（Neottopteris nidus）和地生网脉铁角蕨（Asplenium finlaysonianum）的光合特征和光合诱导特性,来研究不同生态型蕨类植物的光斑利用策略。研究结果表明,2种蕨类植物的最大净光合速率、暗呼吸速率、表观量子效率、光饱和点和光补偿点没有显著差异,但网脉铁角蕨的最大气孔导度远远高于鸟巢蕨,表明后者具有更强的光合水分利用效率。在暗处理3／J＇,时接着光照（光强为20I~mol-m-2,s。‘）30分钟后,网脉铁角蕨的初始气孔导度显著高于鸟巢蕨。连续照射饱和强光后,网脉铁角蕨达到最大净光合速率50％（T50％）和90％的时间（T90%）比鸟巢蕨短：网脉铁角蕨和鸟巢蕨的T50%分别为0．57和5．31分钟,T90％分别为5．85和26．33分钟。诱导过程中,气孔导度对强光的响应明显滞后于净光合速率。鸟巢蕨达到最大气孔导度的时间明显比网脉铁角蕨慢,但在光合诱导消失过程中2种蕨类植物的光合诱导维持能力却没有显著差异。上述结果表明,与大多数地生林下植物（如网脉铁角蕨）相比,附生鸟巢蕨的水分保护比碳获得更重要,但却限制了附生蕨对光斑的利用。