Ribosomal DNA spacer-length variation inSecale spp. (Poaceae)

Variation in ribosomal DNA spacer length was analysed in 23 populations of 12Secale spp. by restriction enzyme analysis. Digestion of rDNA with Taq I endonuclease enzyme yields spacer fragments that include the subrepeat array and the non-repetitive region downstream of the array. Extensive spacer length variation existed in most species with Taq I fragment lengths ranging from 0.9–3.1 kb. These length variants have been attributed to the differences in number of 134 bp spacer subrepeats within rDNA arrays.S. silvestre was the only species to exhibit a unique spacer length variant of 0.9 kb and this was shown to result from the presence of an extra Taq I site in the spacer. rDNA spacer length frequencies were determined for the species. These frequencies were used to derive phenetic relationships between the species by numerical taxonomic methods. In plots constructed fromGower's distance matrices,S. silvestre appeared well separated from the major cluster consisting of the other species. On the basis of morphological and cytogenetic criteria,S. silvestre is considered the most ancient species. The rDNA data is consistent with this interpretation as it shows a clear differentiation ofS. silvestre from all the other species based on length and nucleotide sequence composition of the spacer region.     

The inheritance of rye seed peroxidases

Summary Genetic analyses were conducted on peroxidase of the embryo and endosperm of seeds of one open pollinated and six inbred lines of cultivated rye (Secale cereale L.), and one line of Secale vavilovii Grossh. The analyses of the individual parts of the S. cereale seed yield a total of 14 peroxidase isozymes. Isozymes m, a, b, c, d, e, f and g (in order from faster to slower migration) were found in the embryo plus scutellum, while isozymes 1, 2, 3, 4, 5 and 6 (also from faster to slower migration) were peculiar of the endosperm. S. vavilovii has isozymes m, c₁, d, e, f and g in its embryo plus scutellum, and isozyme 2 in the endosperm. Segregation data indicated that at least 13 different loci would be controlling the peroxidase of S. cereale. Isozymes a and b are controlled by alleles of the same locus, all the other loci have one active and dominant allele coding for one isozyme, and other null and recessive allele. The estimation of linkage relationships shows that five endosperm loci are linked, and tentative maps are shown. A possible dosage effect and the existence of controlling gene(s) for endosperm isozyme 4 is reported. All these data and the high frequency of null alleles found are discussed in relation to recent reports.     

Evolution and systematic relationships in theTriticeae (Poaceae)

The evolution and taxonomic relationships in theTriticeae are discussed with the view to highlight aspects of this agronomically important group of plants, which may be of interest to molecular biology. Some of these aspects are addressed in more detail in adjoining papers in which specific genomic loci have been examined at the DNA sequence or isozyme level. Aspects discussed include the systematics and geographic distribution of theTriticeae species, isozyme and chromosome pairing studies on some of the species as well as more recent developments in DNA analyses. A survey of the systematics of theTriticeae indicated that the genomic system ofLöve is probably the most useful starting point for interpreting molecular data even though the system has many problems from a taxonomic point of view. The geographical distribution ofTriticeae species, using both published and unpublished data, suggested that information of this type taken together with the theory of continental drift provides a broad time-span for considering data from DNA sequence studies. The significance, and modes of analyses, of isozyme studies were assessed because they often provide valuable characters in determining relationships between species. The main character underlyingLöve's andDewey's analyses of theTriticeae, namely chromosome pairing, is discussed with particular reference to isozyme studies to show that in some cases, such as species ofHordeum sensu lato, consistent relationships are obtained. Finally, new developments in understanding chromosome structure are considered in relation to the above variables in the taxonomy and evolution of theTriticeae.     

Esterase isozyme variation in theEragrostis curvula complex (Poaceae)

The biosystematic relationships of the apomictic complexEragrostis curvula s. lato, is investigated by disc electrophoresis of seed extracts to obtain esterase patterns of 23 accessions representing the morphological variants of this complex: curvula, conferta, robusta, chloromelas and lehmanniana. The zymograms thus obtained were classified into four groups on the basis of the presence of certain bands taken as characteristic and constant markers. Within each group variations were found in strict accordance with the morphological and cytogenetic data available on the complex. Cluster analysis showed similarity levels between the strains studied, representing different genomic groups. The esterase pattern proved useful as an additional criterion for identifying the individual taxa making up the complex and for evaluating their reciprocal relationships.     

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.     

Subgroups in theHordeum patagonicum complex (Poaceae)

Seeds of theHordeum patagonicum complex were collected from the field and grown in the greenhouse. The aim was to take a sample of members of the complex, and on the basis of the phenotypic similarities in some morphological and physiological characters, determine whether distinct groups exist. When cluster analyses, to generate hypotheses, and orthodox statistical procedures, for hypotheses obtained a priori, were applied to the reproductive morphology, germination and flowering patterns, onlyH. patagonicum subsp.magellanicum, out of the five recognized taxa, could be distinguished consistently. The remaining four taxa, which overlapped considerably, could be re-formed into three groups whose centroids were different but whose ranges of variation were not distinct from each other. We conclude that the highly cross-compatible members of theH. patagonicum complex, first defined as species and later redefined as subspecies are probably no more than biotypes.     

Species relationships inFestuca (sect.Ovinae,Poaceae)

Morphological and seed protein studies of selected species ofFestuca were performed to elucidate the relationships between species of sect.Ovinae and their affinity with other fescues.Festuca rubra andF. heterophylla (extravaginale group) had higher phenetic affinity with taxa of sect.Scariosae, Montanae andBovinae than with members of their own section. 5 species of the intravaginale group (F. ovina, F. capillata, F. valesiaca, F. sulcata, F. trachyphylla) were fairly uniform and well defined by protein and morphological data. This supports current systematic thinking and does not agree withHackel's (1882) classification of this group.     

Gene flow inAgrostis tenuis (Poaceae)

The copper tolerance of both adult plants and their seedlings, ofAgrostis tenuis from sites within and outside, but neighbouring the Parys Mountain mine (North Wales) were studied. Intensive sampling was carried out along a line starting from the mine boundaries to a distance of five miles away in the direction of the prevailing wind, and of two miles in an upwind direction. The test have shown that mine populations are very tolerant. In outside populations on soils with reduced of very low copper content the mean tolerance level of the seedlings is considerably higher than that of corresponding adult plants, while both show a gradual decrease in tolerance with distance from the mine. The gene flow moves in the direction of the prevailing wind transfering tolerant characters from the mine to the normal populations. The selective pressures on the toxic soil of the mine are quite strong and permit ± tolerant plants only, while on normal soils these pressures are not so severe, so that populations may be rather heterogenous regarding their tolerance towards copper.     

Intergeneric relationships ofLolium,Festuca, andVulpia (Poaceae) and their phylogeny

Morphological and seed protein analyses of 26 species of the generaLolium, Festuca andVulpia confirmed their close systematic affinities. Six inflorescence characters readily differentiatedFestuca fromLolium. Protein similarities betweenFestuca of sect.Bovinae and cross-pollinated species ofLolium, coupled with cytogenetic and crossability data, substantiate that they should be united into one genus.Vulpia had phenetic similarities with sect.Scariosae, Montanae andOvinae ofFestuca. Lolium, Festuca, andVulpia are most likely derived from a common ancestral form which was close toFestuca pratensis andLolium perenne.     

Sequence variation at theSec-1 locus of rye,Secale cereale (Poaceae)

This paper describes the structure of a 9.2-kb repeat unit of DNA, which represents one-secalin gene and spacer sequence located at theSec-1 locus on the short arm of chromosome 1 of rye. The gene units at theSec-1 locus comprise 1.1 kb representing the gene and 8.1 kb of spacer sequence separating the genes. A sequence comparison of nine genes and their promoter regions from theSec-1 locus, reveals that there is greater variation within the coding sequence than there is within the promoter regions. The gene sequence variation is discussed in terms of the size variation seen for the-secalin proteins in rye species. The results include a comparison of promoter sequences from members of the Triticeae to examine the degree of conservation between other seed storage protein genes.     

The intergeneric hybrid origin ofPoa labradorica (Poaceae)

Phytogeographic, ecological, morphological, anatomical, cytological, isozyme, and endonuclease restriction site evidence is presented demonstrating thatPoa labradorica is of intergeneric hybrid origin betweenDupontia fisheri subsp.psilosantha (pistillate parent) andPoa eminens (staminate parent).Poa labradorica is endemic to recently deglaciated areas of northern Quebec and Labrador which are within the overlapping range of the putative parents. Incapable of sexual reproduction,P. labradorica dominates large areas of coastal marsh through vigorous production of rhizomes. Most morphological, anatomical, cytological, isozyme, and DNA characters studied inP. labradorica showed states intermediate or additive to those of the putative parents.     

Delimitation and classification ofBromus fasciculatus (Poaceae)

The syntypes ofBromus madritensis var.delilei Boiss. comprise two different elements: specimens ofB. haussknechtii Boiss. and ofB. fasciculatus C. Presl s.l. By its lectotypification and on the basis of new morphological characters, the validation ofB. fasciculatus subsp.delilei (Boiss.)H. Scholz 1971 was substantiated. This subspecies represents the eastern marginal segregate of the south-Mediterranean species.B. fasciculatus var.alexandrinus Thell. is a minor variant of the typical subspecies.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

Genome relationship betweenPsathyrostachys huashanica andP. fragilis (Poaceae)

Hybrids between the Chinese endemic speciesPsathyrostachys huashanica Keng and the SW. Asian speciesP. fragilis (Boiss.)Nevski (all 2n = 14) developed normally but were completely sterile. Meiotic analyses revealed a high chiasma frequency indicating that the two species as well asP. juncea (Fisch.)Nevski share the same basic genome (called N). The hybrid nature of the plants was established through karyotype analysis and Giemsa C-banding.     

Genomic relationships within theElymus parviglumis group (Triticeae: Poaceae)

Genomic relationships of 13 tetraploid species within the AsiaticElymus parviglumis group containing the SY genomes were assessed by analysing chromosome pairing at metaphase I of the parental species and their interspecific hybrids. Two major genomic subgroups among the tetraploids were identified from the cluster analysis of the averaged c-values, namely, theE. caucasicus subgroup (two species) and theE. parviglumis subgroup (11 species). The genomic affinity of theElymus species is associated with the interspecific geographic distance.After October 1, 1993     

Relationships betweenOryza species (Poaceae) based on 5S DNA sequences

Relationships between 9Oryza species, covering 6 different genomes, have been studied using hybridization and nucleotide sequence information from the5S Dna locus. Four to five units of the major size class of 5S DNA in each species, 55 units in all, were cloned and sequenced. Both hybridization and sequence data confirmed the basic differences between the A and B, C, D genome species suggested by morphological and cytological data. The 5S DNA units of the A genome species were very similar, as were the ones from the B, C, and D genome-containing species. The 5S DNA ofO. australiensis (E genome) grouped with the B, C, D cluster, while the units ofO. brachyantha (F genome) were quite different and grouped away from all other species. 5S DNA units fromO. minuta, O. latifolia, O. australiensis, andO. brachyantha hybridized strongly, and preferentially, to the genomic DNA from which the units were isolated and hence could be useful as species/genome specific probes. The 5S DNA units fromO. sativa, O. nivara, andO. rufipogon provided A genome-specific probes as they hybridized preferentially to A genome DNA. The units fromO. punctata andO. officinalis displayed weaker preferential hybridization toO. punctata DNA, possibly reflecting their shared genome (C genome).     

Biochemical variation to determine phylogenetic relationships betweenHordeum chilense and other American species of the genusHordeum (Poaceae)

Slab gel electrophoresis techniques have been applied to the study of isozyme and kernel protein patterns in 20 accessions ofHordeum chilense and related species in order to elucidate their phylogenetic relationships. On the basis of our results we can conclude that: (1) Conventional classification based on morphological characters does not totally agree with biochemical data. (2) Sectt.Anisolepis andCritesion seem to be clearly differentiated. (3) The accessions classified asH. compressum present biochemical phenotypes quite different from the rest of the species. (4)H. stenostachys, H. muticum andH. chilense constitute a group of variable species with many biochemical similarities and close phylogenetic relationships. (5) The evolutionary pattern of these American species seems to follow a model of reticulate evolution.     

Phylogenetic relationships of Deschampsia antarctica (Poaceae): Insights from nuclear ribosomal ITS

Deschampsia antarctica E. Desv. is the only monocot in the Antarctic floristic zone. We evaluated the phylogenetic relationships of Deschampsia antarctica to other grasses using parsimony as the optimality criterion. Five different sets of gap, transversion and transitions costs were explored to analyze the effect of parameter choice on the phylogenetic results. Both internal transcribed spacers (ITS1 and ITS2) and the 5.8S subunit of nuclear ribosomal DNA were included in the analysis. A total of 43 species were analyzed including seven species of Deschampsia. Deschampsia antarctica forms a well supported group with five species of Deschampsia. Deschampsia does not appear monophyletic as D. flexuosa (L.) Trin. is not included in this clade. The clade to which D. antarctica belongs is sister to some Aveneae in all analyses. This study is the first contribution that evaluates the phylogenetic position of D. antarctica in relation to other species of Deschampsia.     

Relationships between five species ofLolium (Poaceae)

Morphological analysis and electrophoresis of seed proteins of fiveLolium species disclosed that they form two distinct groups corresponding to those recognized from compatibility data.Lolium temulentum andL. remotum of the self-pollinated group were shown to be distinct but closely related species. Morphological intergradation and high similarities between protein profiles ofL. perenne, L. multiflorum andL. rigidum (cross-pollinated species) suggest little genetic differentiation between these taxa. This implies that treatment at the infraspecific level might better accomodate the data.     

Genome relationships in theElytrigia group of the genusAgropyron (Poaceae) as indicated by seed protein electrophoresis

Agropyron bessarabicum (2n = 14),A. rechingeri (2n = 28),A. junceiforme (2n = 28),A. elongatum (2n = 14),A. flaccidifolium (2n = 28) andA. scirpeum (2n = 28) were studied by isoelectric focusing of seed soluble proteins.—The protein profiles obtained from the six taxa showed a striking degree of similarity; typically they consist of 40 bands. No qualitative but only quantitative differences (in the intensity of some bands) were found.—Combined with the cytological information available these protein data indicate that the two polyploid complexes must be placed in the recently erected genusThinopyrum with the genome designations:T. bessarabicum J^j1 J^j1,T. sartorii (=A. rechingeri) J^j1 J^j1 J^j3 J^j3,T. junceiforme J^j1 J^j1 J^j2 J^j2,T. elongatum J^e1 J^e1,T. flaccidifolium J^e1 J^e1 J^e1 J^e1 andT. scirpeum J^e1 J^e1 J^e2 J^e2.