Isozyme evidence for host races of the fungus Atkinsonella hypoxylon (Clavicipitaceae) infecting the Danthonia (Poaceae) complex in the southern Appalachians

On granite outcrops around Highlands, North Carolina four sympatric Danthonia grass species are infected by the fungus Atkinsonella hypoxylon (Clavicipitaceae). Danthonia epilis and D. sericea (rare hosts) are infected only in this small region, although they range more widely, while D. compressa and D. spicata (widespread hosts) are infected throughout their ranges in eastern North America. To test the hypothesis that infection of the rare hosts has recently spread from the widespread hosts, the genetic structure of fungal populations on different Danthonia hosts was investigated by isozyme electrophoresis. Two hundred isolates were collected from 46 sites and analyzed for ten polymorphic isozyme loci to examine levels of variation, genetic differentiation, and gene flow among fungal populations infecting different hosts. Genetic identities (I) indicated that isolates from within each of the widespread and rare host species pairs were homogeneous (I > 0.99), but the pairs were highly differentiated (0.66 < I < 0.69). With one exception there was no overlap in fungal genotypes isolated from the widespread hosts (12 genotypes) vs. the rare hosts (seven genotypes). Two plants of D. compressa growing sympatrically with infected D. epilis at one site were infected with a genotype characteristic of the latter species. Estimates of gene flow suggest a high degree of host-mediated reproductive isolation in A. hypoxylon, indicating that the fungus forms distinct, long-standing host races or sibling species in the southern Appalachians.     

Intergenomic rearrangements after polyploidization of Kengyilia thoroldiana (Poaceae: Triticeae) affected by environmental factors

Polyploidization is a major evolutionary process. Approximately 70–75% species of Triticeae (Poaceae) are polyploids, involving 23 genomes. To investigate intergenomic rearrangements after polyploidization of Triticeae species and to determine the effects of environmental factors on them, nine populations of a typical polyploid Triticeae species, Kengyilia thoroldiana (Keng) J.L.Yang et al. (2n = 6x = 42, StStPPYY), collected from different environments, were studied using genome in situ hybridization (GISH). We found that intergenomic rearrangements occurred between the relatively large P genome and the small genomes, St (8.15%) and Y (22.22%), in polyploid species via various types of translocations compared to their diploid progenitors. However, no translocation was found between the relatively small St and Y chromosomes. Environmental factors may affect rearrangements among the three genomes. Chromosome translocations were significantly more frequent in populations from cold alpine and grassland environments than in populations from valley and lake-basin habitats (P<0.05). The relationship between types of chromosome translocations and altitude was significant (r = 0.809, P<0.01). Intergenomic rearrangements associated with environmental factors and genetic differentiation of a single basic genome should be considered as equally important genetic processes during species'' ecotype evolution.     

Cytology of a monosomic-trisomic Coix gigantea (Poaceae)

Meiotic behaviour of a monosomic-trisomic (2n–1+1=20) of Coix gigantea (Poaceae) showed that, although there are a few segregational irregularities, it is essentially mostly normal. The plant is healthy and semifertile, and through the formation of viable male and female gametes of variable chromosomal constitution it is expected to produce simple and complex aneuploids in its progeny. The origin is believed to have been through nondisjunction of two bivalents, a large and a small in a diploid.     

The measurement of small-scale environmental heterogeneity using clonal transplants of Anthoxanthum odoratum and Danthonia spicata

Summary To assess the scale of micro-environmental heterogeneity perceived by two co-occurring grass species, Anthoxanthum odoratum and Danthonia spicata, cloned tillers of each species were planted into the natural habitat at a range of spacings (from 2 cm to more than 2 m apart) and measured for survival and fecundity over three years. A. odoratum responded to heterogeneity at a scale of 4–8 cm and at a scale of 2–8 m but not to intermediate scales. D. spicata did not respond significantly to heterogeneity. However one genotype infected with the systemic fungus Atkinsonella hypoxylon showed a large response to heterogeneity at both small and large spatial scales. The results showed that the scale and level of environmental heterogeneity as measured by its fitness impact depends on the species and genotype involved. The results indicate that small scale environmental heterogeneity could play a role in the maintenance of sexual reproduction in A. odoratum.     

Inflorescence development in a new teosinte: Zea nicaraguensis (Poaceae)

Inflorescence development in a newly discovered teosinte, Zea nicaraguensis (Poaceae), from Nicaragua has been investigated using scanning electron microscopy (SEM). The SEM examination revealed that the pattern of both male and female inflorescence development was similar to previously described inflorescence in other Zea taxa. Branch primordia were initiated acropetally in a distichous pattern along the rachis of male and female inflorescences. Spikelet pair primordia bifurcated into pedicellate and sessile spikelet primordia. Predictably, pedicellate spikelet development was arrested and aborted in the female teosinte inflorescence. Organogenesis of functional spikelets and florets was similar to that previously described in maize and teosintes. The results were consistent with our hypothesis that both femininity and masculinity share a common mechanism of inflorescence development in Zea and Tripsacum and are in accord with a putative common mechanism of sex determination in the Andropogoneae. Interestingly, this population of teosinte, unique in its ability to grow in water-logged soils, showed a stable pattern of early inflorescence development. Our results also revealed the uncharacteristic presence of inflorescence polystichy in this population of Zea nicaraguensis. We propose this novel phenotypic variation raises the possibility that a domestic evolution of polystichy in maize was enabled by an occasional polystichous phenotypic in teosinte.     

Inflorescence development in a perennial teosinte: Zea perennis (Poaceae)

The ontogeny of tassels and ears in a perennial Mexican teosinte, Zea perennis (Hitchc.) Reeves and Mangelsdorf, was examined using scanning electron microscopy and light microscopy. Ear development follows a pattern previously described for two annual teosintes, Z. mays subsp. mexicana and Z. mays subsp. parviglumis var. parviglumis (race Balsas), and for the bisexual mixed inflorescence in a diploperennial teosinte, Z. diploperennis; it differs from that described for the ear of Z. diploperennis plants grown at the latitudes of Iowa and Wisconsin. Common bud primordia of the ear are initiated in the axil of distichously arranged bracts along the ear axis. These common primordia bifurcate to form paired pedicellate and sessile spikelet primordia. Development of the pedicellate spikelets in the ear is arrested leaving the sessile spikelets, along with the adjoining rachis segment, to form solitary grains enclosed within cupulate fruitcases. The organogenesis of the central spike of the tassel is similar to that previously described in other Zea taxa. This developmental study supports the hypothesis that both femaleness and maleness are derived from and expressed on a common background; it is consistent with the view that the maize ear was derived from the central spike of a male inflorescence terminating a primary branch of the main axis of the inflorescence.     

Calamagrostis hakonensis (Poaceae): Distribution and differentiation of cytotypes

Chromosome counts were made for a total of 540 collections ofCalamagrostis hakonensis from 66 localities ranging from Kyushu to Hokkaido. Distribution and habitat preferences of the cytotypes involved are described. Sympatric occurrence of tetraploids (=semidiploids, 2n=28) and higher polyploids, as well as that of septaploids and plants at hexaploid and/or octoploid levels, was confirmed in several localities, and mixtures of hexaploids and octoploids within a population were frequently observed. Plants at hexaploid and octoploid levels were the most abundant and widespread. An examination of pollen of the voucher specimens showed that tetraploids (amphimictic) had good pollen, while higher polyploids (apomictic) were generally devoid of pollen and very rarely produced moderately good pollen, the grains of higher polyploids being larger than those of tetraploids. Distributions of tetraploids and higher polyploids were more precisely delineated through an examination of pollen of many herbarium specimens. In relation to the processes by which the complicated internal structure ofC. hakonensis has been established, the following subjects are discussed: infraspecific hybridization between ecotypically differentiated populations, enrichment of variability through occasional sexual reproduction expected in plants with more than 2n=42, persistence of the variants by apomictic reproduction, and probable roles of some extinct taxa.     

窄颖仲彬草生物系统学研究

窄颖仲彬草Ｋｅｎｇｙｉｌｉａ ｓｔｅｎａｃｈｙｒａ（Ｋｅｎｇ） Ｊ．Ｌ． Ｙａｎｇ ,Ｙｅｎ ｅｔ Ｂａｕｍ 是分布于我国西部的一种多年生六倍体植物。将其与犬草Ｅｌｙｍｕｓ ｃａｎｉｎｕｓ（Ｌ．） Ｌ．, 鹅观草Ｒｏｅｇｎｅｒｉａ ｋａｍｏｊｉ Ｏｈｗｉ, 糙毛仲彬草Ｋ．ｈｉｒｓｕｔａ （Ｋｅｎｇ） Ｊ．Ｌ．Ｙａｎｇ,Ｙｅｎ ｅｔ Ｂａｕｍ ３ 个种进行了杂交。对亲本及杂种Ｆ１ 代花粉母细胞减数分裂中期Ｉ染色体配对行为进行了观察。减数分裂平均构型分别为： Ｅ． ｃａｎｉｎｕｓ×Ｋ． ｓｔｅｎａｃｈｙｒａ２３－７９ Ⅰ＋ ５－２０ Ⅱ＋ ０－２７Ⅲ; Ｒ．ｋａｍｏｊｉ ×Ｋ．ｓｔｅｎａｃｈｙｒａ１８－２３ Ⅰ＋ １１－６８ Ⅱ＋ ０－０６ Ⅲ＋ ０－０６ Ⅳ; Ｋ．ｈｉｒｓｕｔａ ×Ｋ．ｓｔｅｎａｃｈｙｒａ４－８３Ⅰ＋ １７－３１ Ⅱ＋ ０－５５ Ⅲ＋ ０－２０ Ⅳ＋ ０－０２ Ⅴ。根据以上结果, 结合种的形态特征, 窄颖仲彬草应从鹅观草属Ｒｏｅｇｎｅｒｉａ Ｃ． Ｋｏｃｈ 拟冰草组ＰａｒａｇｒｏｐｙｒｏｎＫｅｎｇ 中组合到仲彬草属Ｋｅｎｇｙｉｌｉａ Ｙｅｎ ｅｔ Ｙａｎｇ。     

New species of cryptochloa and piresia (Poaceae: Bambusoideae)

Two new species are added toCryptochloa,C. unispiculata from Peru andC. capillata from Brazil, the latter transferred from the genusOlyra. The genusPiresia is also augmented by two new species,P. macrophylla from Peru andP. leptophylla from Bahia, Brazil. These species represent, respectively, the westernmost and easternmost limits of distribution of the genus.     

Interspecific crosses inHordeum (Poaceae)

A crossing programme including 30 species and 40 cytotypes within the genusHordeum was undertaken. Viable hybrids were obtained in 302 combinations, 15 of which were intraspecific. Differences in seed set and in germination were observed in crosses between different groups of species. Obtaining crosses between different taxonomic groups was generally more difficult when diploid material was used. Some species, e.g.,H. lechleri, H. jubatum, andH. brachyantherum showed a higher crossability than others. The chromosome numbers of the hybrids were usually those expected from the parental numbers but aneuploid series around the expected numbers were rather frequent. Three cases of unreduced gametes were found. Selective chromosome elimination was restricted to combinations including eitherH. vulgare orH. bulbosum.—Despite a very diverse morphology, all South American diploid species together with the two North American diploidsH. intercedens andH. pusillum appear to be closely related. The hexaploid American speciesH. procerum, H. lechleri, andH. arizonicum are also related. The two North American tetraploid speciesH. jubatum andH. brachyantherum sometimes form semifertile hybrids. The Asiatic speciesH. roshevitzii appears to be related to both North and South American taxa.     

Tripsacum dactyloides (Poaceae): a natural model system to study parthenogenesis

Diplosporous apomeiosis, formation of unreduced embryo sacs primarily of the Antennaria type, followed by parthenogenetic embryo development and pseudogamy (fertilization of the central cell) describe gametophytic apomixis within the Tripsacum agamic complex. Tripsacum dactyloides (Eastern gamagrass) is a close relative of domesticated maize and was chosen as a natural model system to investigate gene expression patterns associated with parthenogenesis. The genome size of diploid sexual and polyploid apomictic T. dactyloides was estimated by flow cytometry to be 7.37 pg (2C), 14.74 pg (4C) and 22.39 pg (6C), respectively. The diploid genome size is thus approximately 1.352 larger than that of maize. The apomeiotic-pseudogamous pathway of seed formation was demonstrated at a rate of 92% by the flow cytometric seed screen (FCSS) with single mature seeds in tetraploid accessions. This number includes twin embryos which were detected in 13% of the seeds analyzed. Fertilization of unreduced egg cells (B_III hybrids) was measured in 10% of apomictic seeds. Autonomous (fertilization-independent) embryo development and fertilization-dependent endosperm formation were confirmed by pollination of tetraploid T. dactyloides with a diploid transgenic maize line carrying an actin::#-glucuronidase (GUS) reporter construct. GUS expression was detected after pollination in the developing endosperm, but not in the embryo. In similar intraspecific crossing experiments with maize, GUS expression was detected in both the embryo and endosperm. A protocol was established for microdissection of embryo sacs and early parthenogenetic embryos of T. dactyloides. Together, these techniques provide new tools for future studies aimed at comparing gene expression patterns between sexual maize and sexual or apomictic T. dactyloides.     

The genus Aciachne (Poaceae)

A lectotype has been selected for Aciachne pulvinata. The name A. uniflora is rejected as nom. illeg. Two new species, A. acicularis and A. flagellifera are described and a key is provided. Leaf anatomy is described and discussed, several diagnostic-characters have been found. Distribution maps and altitudinal ranges are discussed.     

Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient

An invasive variety of Phragmites australis (Poaceae, common reed), the M haplotype, has been implicated in the spread of this species into North American salt marshes that are normally dominated by the salt marsh grass Spartina alterniflora (Poaceae, smooth cordgrass). In some European marshes, on the other hand, Spartina spp. derived from S. alterniflora have spread into brackish P. australis marshes. In both cases, the non-native grass is thought to degrade the habitat value of the marsh for wildlife, and it is important to understand the physiological processes that lead to these species replacements. We compared the growth, salt tolerance, and osmotic adjustment of M haplotype P. australis and S. alterniflora along a salinity gradient in greenhouse experiments. Spartina alterniflora produced new biomass up to 0.6 M NaCl, whereas P. australis did not grow well above 0.2 M NaCl. The greater salt tolerance of S. alterniflora compared with P. australis was due to its ability to use Na(+) for osmotic adjustment in the shoots. On the other hand, at low salinities P. australis produced more shoots per gram of rhizome tissue than did S. alterniflora. This study illustrates how ecophysiological differences can shift the competitive advantage from one species to another along a stress gradient. Phragmites australis is spreading into North American coastal marshes that are experiencing reduced salinities, while Spartina spp. are spreading into northern European brackish marshes that are experiencing increased salinities as land use patterns change on the two continents.     

Multiple homoplasious insertions and deletions of a Triticeae (Poaceae) DNA transposon: a phylogenetic perspective

Background  

Stowaway elements are short, non-autonomous DNA transposons categorized as miniature inverted-repeat transposable elements (MITEs). The high MITE copy number in grass genomes suggests an active history of amplification and insertion, but ongoing MITE activity has only rarely been seen, and ongoing Stowaway activity has never been observed. Thus, a phylogenetic perspective on presence vs. absence of elements in an aligned data set can provide valuable historical insights into the dynamics of MITE acquisition and loss.     

Nomenclatural changes in schizachyrium (Poaceae)

Two morphological variants ofSchizachyrium hirtiflorum are reduced to varietal rank underS. sanguineum. Schizachyrium semiberbe is included as a synonym of the typical variety ofS. sanguineum.     

Aneuraceae (Metzgeriales) and tulasnelloid fungi (Basidiomycota): a model for early steps in fungal symbiosis

A total of 35 population samples of the liverwort genera Aneura (A. pinguis) and Riccardia (R.?latifrons, R. multifida, and R. palmata) were sampled from diverse habitats and geographical provenances in Germany, Austria, and Switzerland. Light and transmission electron microscopy were used to characterise the morphological features of the associations, and phylogenetic analyses based on internal transcribed spacers (ITS) and the D1/D2 regions of the fungal 28S rDNA were used to address diversity and phylogenetic relationships. By comparing the cellular structures of the plant-fungus interactions, we recognised the following states of fungal colonisation within the thalli: fungus-free, epiphytic, intercellular, and intracellular. Colonising hyphae showed dolipores with imperforate parenthesomes, slime bodies, and multilayered walls. Colonised liverwort cells had pleomorphic nuclei and elongated starch-free chloroplasts with distinctive grana. Our analyses revealed six phylogenetic groups of tulasnelloid fungi associated with liverworts, where major lineages mostly share similar host and/or ecological specialisations. The mode of colonisation of the tulasnelloid mycobionts in Aneura and Riccardia sharing identical fungal sequences is different. Consequently, the mode of colonisation may be host-dependent. Finally, our findings demonstrate that Aneuraceae are model organisms for evolutionary studies of symbiotic associations between liverworts and fungi.     

Slipped-strand mispairing in a plastid gene: rpoC2 in grasses (Poaceae)

An exception to the generally conservative nature of plastid gene evolution is the gene coding for the beta" subunit of RNA polymerase, rpoC2. Previous work by others has shown that maize and rice have an insertion in the coding region of rpoC2, relative to spinach and tobacco. To assess the distribution of this extra coding sequence, we surveyed a broad phylogenetic sample comprising 55 species from 17 angiosperm families by using Southern hybridization. The extra coding sequence is restricted to the grasses (Poaceae). DNA sequence analysis of 11 species from all five subfamilies within the grass family demonstrates that the extra sequence in the coding region of rpoC2 is a repetitive array that exhibits more than a twofold increase in nucleotide substitution, as well as a large number of insertion/deletion events, relative to the adjacent flanking sequences. The structure of the array suggests that slipped-strand mispairing causes the repeated motifs and adds to the mechanisms through which the coding sequence of plastid genes are known to evolve. Phylogenetic analyses based on the sequence data from grass species support several relationships previously suggested by morphological work, but they are ambiguous about broad relationships within the family.      

Phylogenetic relationships inSecale (Poaceae): An isozymatic study

The genetic frequencies of 9 isozyme loci have been estimated in 23 samples of 4 species ofSecale by means of starch gel electrophoresis. The populations ofS. silvestre andS. vavilovii were monomorphic and uniform within each species, those ofS. montanum andS. cereale were polymorphic for most of the isozyme loci. On the basis of isozyme patterns as well as allelic and genotypic frequencies of isozyme loci,S. silvestre can be distinguished fromS. vavilovii, and both fromS. cereale andS. montanum; but there is no clear differentiation between the two latter species. Clusters constructed from genetic distances separateS. silvestre andS. vavilovii, whereasS. cereale andS. montanum were grouped together. The isozymatic data presented here, along with cytogenetic and life habit data, agree with the generally admitted existence of 4 species inSecale, and support the relationships suggested byKhush & Stebbins (1961).     

Intergeneric hybridizations with Eremopyrum (Poaceae)

Intergeneric crosses were made between the four species of Eremopyrum (2n = 14, 28) and 16 species (2n = 14, 28, 42) from 8 genera of the Triticeae. Seed set was obtained in 22 of 42 different combinations. Only 9 resulted in progeny and only one was diploid. The hybrids were morphologically intermediate between the parents and it has been confirmed that perenniality dominates over annuality. The very low pollen fertility of the hybrids was caused by very low chromosome pairing in meiosis and supports the uniqueness of the Eremopyrum genome. Autosyndesis between the chromosomes of E. orientale is proposed and for that reason reevaluation of the relationships within Eremopyrum is needed.