Genetic determination of pappus part number in the annual hybridMicroseris B 87 (Asteraceae—Lactuceae)

The ChileanMicroseris pygmaea has a ten-part paleaceous pappus while the CalifornianM. bigelovii has five pappus parts on each achene. Hybrids between the two species have between five and ten pappus parts with averages below 7.5. Hybrid B 87 has an F 1 average value of 6.7 pappus parts. 140 F 2 plants were raised from this hybrid, and 12 F 3 families were obtained by selfing from F 2 plants. One larger F 4 family has been raised. Pappus part number in all of these is still canalized between 5 and 10. Variation within these limits is genetically determined by a quantitatively acting polygenic system. Modeling of this system suggests that a minimum of four, but probably not many more, genes are involved. This opens the possibility of a complete genetic analysis of the system.     

Four additive genes determining pappus part numbers inMicroseris annual hybrid C34 (Asteraceae/Lactuceae)

Microseris strain C34 is a hybrid between the Chilean speciesM. pygmaea (10 pappus parts) and the CalifornianM. bigelovii (5 pappus parts). The F1 specimen had from 5 to 10 pappus parts per achene with an average of 6. F2, F3 and F4 plants derived from this hybrid by spontaneous selfing show segregation for the average number of pappus parts. Four segregating unlinked genes could be demonstrated, each with an allele determining 5 pappus parts from thebigelovii parent, one with an allele determining 10 pappus parts, three with null alleles from thepygmaea parent. The expected average pappus part number is the arithmetic mean of the 5- and 10-determining factors. Considerable environmental and developmental influences, both random noise and systematic shifts, could be demonstrated to influence the phenotypic expression. The parallel hybrid strain B87 has two 10-alleles rather than one in itspygmaea genome. The evolution of the pappus part genes ofM. pygmaea from those of abigelovii-like ancestor seems to demand the concerted (non-independent) mutation of at least two genes.     

Variability of the inflorescence ofMicroseris laciniata (Compositae: Cichorieae)

Quantitative characters of the flowering head of a garden population ofMicroseris laciniata were scored during the second, third, and fourth season of growth. Number of achenes per head, number of phyllaries per head and the average number of pappus parts per achene in single heads show significant plant to plant variation. Achenes per head and pappus parts per achene were scored in identical plants in two subsequent seasons. The number of pappus parts per achene varies freely between five and ten. This contrasts with annual species ofMicroseris in which either five or ten pappus parts are found, depending on the species. In spite of a clear plant-specific average of pappus parts, both high and low pappus part determination can be demonstrated in all specimens. The number of pappus parts depends on the position of an achene on the receptacle, marginal achenes usually having fewer pappus parts than central ones. This gradient is not closely correlated with the position of an achene on the genetic spiral.     

Single-gene heterozygotes derived from the polygenic pappus part system ofMicroseris hybrid C34 (Asteraceae—Lactuceae)

Microseris strain C34 is a hybrid between the Chilean speciesM. pygmaea (10 pappus parts) and the CalifornianM. bigelovii (5 pappus parts). The plants are propagated by selfing from the original hybrid specimen. Each plant has from 5 to 10 pappus parts per achene with an average value that is additively determined by four unlinked quantitatively acting genes. Single-gene heterozygote sublines have been obtained for two of these,pp-1 (shown to be linked to a modifier of acid phosphatase-1) andpp-4. Sublines homozygous for all four additive genes show residual genetic variation influencing pappus part number. At least one additional gene can be demonstrated by its linkage with leucine aminopeptidase-1. Lines for the further characterization of these hypostatic genes are selected.     

A second marker enzyme in the genetics of pappus part numbers inMicroseris hybrid B87 (Asteraceae,Lactuceae)

CrossingMicroseris pygmaea (10 pappus parts) withM. bigelovii (5 pappus parts) results in hybrids with variable pappus part numbers between 5 and 10. Previous work has shown that a system of four additively acting genes determines the average pappus part numbers of these hybrids. In hybrid B87 two genes have a 10-determining and a 5-determining allele each, two others a 5-determining and a null (inactive or missing) allele. Genetic linkage of one of the latter with the enzyme geneEsterase-1 and the leaf shape genespatulate leaves has been demonstrated. Here we demonstrate linkage between one of the two 10-determining genes and the enzyme locusEsterase- Y/B. The genotypes in the pappus part system of many specimens can now be fully determined. This is a major advance for the analysis of the evolution of this additive polygenic system.Genetics of Pappus Part Numbers inMicroseris Hybrid B87, II.—Part I:Bachmann & al. 1981.     

Genetic components of heterocarpy inMicroseris hybrid B87 (Asteraceae,Lactuceae)

Microseris B87 is derived from a single hybrid specimen betweenM. pygmaea with few, weakly hairy peripheral achenes and aM. bigelovii with many, strongly hairy peripheral achenes. Offspring through the F4 and F5 generations obtained by spontaneous selfing were analyzed for the segregation of quantitative and qualitative characters relating to achene dimorphism. The phenotypic effects of two previously identified unlinked genes determining the relative number of outer achenes are characterized in partially and completely homozygous sublines. We show that two morphological markers genetically linked to one of these genes are themselves regulated by the system inducing heterocarpy. Not more than two more unlinked genes are involved in the genetic basis of the heterocarpic response. The interaction of these genes in determining the heterocarpy phenotypes is discussed in the framework of a model postulating genes for a morphogen gradient across the capitulum and genes responding to this gradient.     

Genetic and Non-genetic Variation in the Number of Pappus Parts in Microseris douglasii Strain D37 (Asteraceae, Lactuceae)

The pappus of the annual species of Microseris usually consistsof a constant number of five paleaceous, awn-tipped scales.In some populations of Microseris douglasii, plants with variablenumbers of pappus parts, always five or fewer, are frequent.Here we study inbred lines derived from a plant with an averageof slightly less than five pappus parts (strain D40), from aplant with an average near three pappus parts (strain B14),and from F2 plants of a hybrid (D37) between these two strains.The average deviation from five pappus parts is subject to varioussources of non-genetic variation. The peripheral achenes ineach head have more pappus parts than the central ones, andthe overall average number decreases in later-formed capitulaof a plant. Both effects are strongly affected by environmentalfactors. Against this background, we show (polygenic) geneticvariation for the average deviation from five, and additionalheritable differences in the strength of the gradient in pappuspart numbers from the periphery to the centre of the capitulum,while the decrease of pappus part numbers in later-formed capitulais essentially identical in both parental strains, even in itsresponse to environmental factors. Visible phenotypic variationdue to all sources decreases as the average nears a constantnumber of five pappus parts Asteraceae, canalized characters, developmental genetics, inflorescence, Lactuceae, Microseris douglasii, phenotypic plasticity     

Comparative capitular morphology and anatomy of Coreopsis L. and bidens L. (compositae), including a review of generic boundaries

The capitular and floral morphology and anatomy ofBidens L. andCoreopsis L. were studied. All the North American species ofCoreopsis were studied. Selected species ofBidens from North and South America andCoreopsis from South America were included. The results were compared with previous observations on African species ofBidens (incl.Coreopsis). Emphasis was given to character states of the ray florets, paleae, stylearm apices, outer phyllaries, achenes, and pollen grains. Some of the character states are unique features ofCoreopsis, e.g., globular and elongately conical receptacles, deltoid outer phyllaries, truncate and indistinctly 3–5-dentate, 3–4-lobed ray florets, narrowly spathulate paleae, subulate paleae with linear-filiform upper half, hairy and apically 3-cleft paleae, truncate, convex or shallowly conical stylearm apices with the sweeping hairs limited to the area above the stigmatic surfaces and the orbicular to circular achenes. The cylindric setaceous pappus bristles so commonly encountered inBidens are unknown inCoreopsis. The pappus bristles inCoreopsis are paleaceous but similar, though thicker ones are also found in African species ofBidens (incl.Coreopsis) with winged achenes. Twin-celled hairs (setulae) with differing degrees of wall thickness are found on the achenes ofCoreopsis sect.Pseudoagarista (Mexico and South America),Coreopsis sect.Pugiopappus (California), AfricanBidens with winged achenes (e.g.,B. prestinaria, B. macroptera) and some North AmericanBidens (e.g.,B. aristosa). Similar sclerotic parenchyma make up the achenial wings of species in both genera. These may be interpreted as homologous structures, indicating the underlying similarity of these taxa and their derivation from a common ancestral stock.     

Intraspecific evolution ofMicroseris pygmaea (Asteraceae,Lactuceae) analyzed by cosegregation of phenotypic characters (QTLs) and molecular markers (RAPDs)

The Chilean annual,Microseris pygmaea, has differentiated in distinct coastal and inland series of populations after long-distance dispersal from western North America. Two plants from the most diverse biotypes were crossed, a large F₂ was raised and analysed for segregation of 30 phenotypic characters. Segregation of molecular markers (47 RAPDs, 1 RFLP, 2 isozymes) was determined in a subpopulation of 45 plants which include all extremes for the phenotypic characters. 32 marker/character cosegregations were significant at the 1% level in t-tests between dominant and homozygous recessive marker genotypes. Considering linkage among markers and pleiotropy of certain marker loci, the number of independent quantitative trait loci (QTLs) is reduced to about 18. Interactions among 2 or 3 QTLs affecting one character have been characterized. The phenotypic differentiation ofM. pygmaea during its evolution from a single founder individual begins to be understood at the level of single-gene mutants.     

Sexual tetraploid and apomictic pentaploid populations ofHieracium pilosella (Compositae)

Five species are recognized inHieracium subgen.Pilosella sect.Pilosellina Fries. Four are diploid (2x, 2n = 18), one (H. pilosella L.) is highly variable morphologically and cytologically (from 2x to 10x), in its mode of reproduction (self-incompatibility, agamospermy, amphimixis, apo-amphimixis) and in its hybridization pattern. A part of this huge agamic complex was analysed by comparing sexual 4x and apomictic 5x plants (crossing and germination experiments, measurements of vegetative reproduction by stolons etc.). In the experimental garden apomictic 5x produced more stolons than the sexual 4x plants and the total length of the stolons per rosette was greater. However, in nature, the competitive potential of the sexual plants seems to be higher, presumably as a result of the higher mortality of ramets in 5x. Sexual 4x plants often grow in dense and grazed grass vegetation, whereas 5x apomicts often occur in dunes with patchy vegetation. Apomicts produce more capitula per rosette, and sexual rosettes form only about 60% of the number of viable achenes as compared to apomictic ones. Therefore, apomicts appear to be characterized by a greater colonizing ability than sexual plants. Apomictic plants produce equal numbers of viable achenes under conditions of both open pollination and isolation. Sexual plants do not form any viable achenes after isolation and produce a somewhat lower percentage of achenes after open pollination than do apomictics. 5xreproduce exclusively apomictically. Apo-amphimixis was never observed in pentaploids and only very rarely in tetraploids. Addition hybrids are very rare. The cross sexual 4x × apomictic 5x failed in 70% of the attempts, but the recombination of genomes carrying genes for apomixis is possible and results in apomictic 4x and sexual 5x, both with a reduced number of viable achenes. In nature sexual and apomictic plants may occur in close proximity. In such cases the germination rate of the achenes of 4x and 5x is lower; this may indicate that apomictic plants fertilize sexual plants in nature (unidirectional gene-flow). 5x plants form euploid gametes carrying two or three genomes. The results of the crossing experiments can be explained in terms ofNogler's theory of monogenic inheritance of apospory.Variation and evolution inHieracium subg.Pilosella sect.Pilosellina I.     

Lipochaeta and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): establishing their natural limits and a synopsis

We restrict the genusLipochaeta to the allopolyploid species of the typical section.Lipochaeta s.str. is interpreted to be the result of an intergeneric hybridization betweenMelanthera and a presently unknown taxon, perhaps of the genusWedelia. Lipochaeta is characterized, in addition to its allopolyploidy (n=26), by having both flavonols and flavones, disk corollas with 4 lobes, achenes tuberculate at maturity, the disk achenes flattened to slightly biconvex, and ray achenes obcompressed.Lipochaeta sect.Aphanopappus andWollastonia are here reduced to synonymy underMelanthera. We transfer 14 HawaiianLipochaeta and one New Caledonian species as well as the AsianWedelia prostrata toMelanthera. These transfers, along with the species in Africa and North America, bring the number of species in the genus to 35.Melanthera is delimited by an abruptly narrowed to truncate and flattened top of the achene, (0−)1–15(−20) often unequal, ciliate or barbellate, caducous pappus bristles immediately surrounding the corolla, involucral bracts and receptacular paleae with many veins forming longitudinal striations, and n=15. The florets are 5-merous, the corollas are yellow or white, and rays are absent (in white-flowered species) or present and neutral or fertile. In dealing with species formerly placed inLipochaeta, the GalapagosL. laricifolia is here transferred from the illegitimate generic nameMacraea toTrigonopterum and the BrazilianL. goyazensis is transferred toAngelphytum. We maintain the earlier reduction ofEchinocephalum underMelanthera and reduce all three taxa originally described in it to one,M. latifolia.     

Systematic overview of Krigia (Asteraceae-Lactuceae)

Cytological and morphological variation among Krigia species is examined. Krigia exhibits a broad range of chromosome numbers including n = 4, 5, 6, 9, 10, 15, and 30. Section Krigia is characterized by reflexed phyllaries and a base chromosome number of x = 5, while section Cymbia is characterized by erect phyllaries and chromosome numbers of n = 4, 6, and 9. The micromorphological characteristics of achenes, pappus, styles, corolla, pollen, stomata, and trichomes are documented using scanning electron microscopy. Among these, the pappus shows the greatest diversity and three major types are identified: 1) a pappus of many bristles and scales, as in K. dandelion, K. montana, and K. biflora; 2) a pappus of five bristles and five scales, as in K. virginica and K. occidentalis; and 3) an absent or highly reduced pappus, as in the K. cespitosa complex and K. wrightii. Thirty-five cytological and morphological characters are subjected to phylogenetic analyses. The two sections, Krigia and Cymbia, form monophyletic lineages. Within section Krigia, the annual species, K. virginica, forms an independent clade, while the perennial species, K. dandelion, K. biflora, and K. montana, form a monophyletic clade. Krigia montana and K. biflora are identified as sister species and a hybrid between these has been identified. The hybrid is more similar morphologically to K. montana than K. biflora. Within section Cymbia, phylogenetic relationships among K. wrightii, K. occidentalis, and K. cespitosa are uncertain. Nine taxa of Krigia are herein recognized: K. dandelion, K. biflora, K. biflora var. viridis (comb. nov.), K. montana, K. virginica, K. wrightii (comb. nov.), K. occidentalis, K. cespitosa, and K. cespitosa f. gracilis (comb. nov.). Phylogenetic relationships among 12 taxa of Krigia species are compared using various combinations of morphology, chloroplast DNA, and nuclear ribosomal DNA data. Tree topologies from different combinations of data are largely congruent. The most resolved phylogenetic tree is obtained using the combined data from morphology, chloroplast DNA, and nuclear ribosomal DNA.     

一年生短命植物疏齿千里光果实异形性的生态学意义

疏齿千里光(Senecio subdentatus)是分布在新疆北部古尔班通古特沙漠中的一种具异形瘦果的菊科一年生短命植物。将野外观测与室内实验相结合, 对该物种异形瘦果的形态、扩散和萌发特性, 以及异形瘦果产量与植株大小的关系进行了研究, 并对其生态学意义进行了探讨。结果表明: 疏齿千里光果序中的外围果和中央果均为柱形, 但前者为淡黄色, 后者为褐色, 且二者在大小、冠毛长度及果皮微形态等方面均存在明显差异。两种瘦果均以单个果实为扩散单元, 且在静止空气中的降落速度和在1 m·s^-1与2 m·s^-1风速下的扩散距离无显著差异, 说明虽然果实大小和冠毛长度对瘦果扩散具有不同的影响, 但对其整体扩散能力无明显影响。在各温变周期(5/2、15/2、20/10、25/15和30/15 ℃)处理中, 淡黄色外围果的萌发率均高于褐色中央果的, 且不同温度间两种瘦果的萌发率均存在显著差异, 但光照条件对其无显著影响。果序中的中央瘦果数明显多于外围果的, 且植株中外围果所占比例与植株大小间呈显著负相关关系, 而中央果所占比例与植株大小间呈显著正相关关系。这些特点说明, 该物种的小植株倾向于产生较多较易萌发的外围果, 大植株倾向于产生较多不易萌发的中央果。在古尔班通古特沙漠不可预测的极端环境中, 疏齿千里光可通过异形瘦果间的萌发差异及调节其不同大小植株中异形瘦果的比例, 来减少同胞后代之间的竞争, 增加其对不同微环境条件的生态适应性。     

青藏高原高寒草甸两种优势植物的生长及其CNP化学计量特征对模拟增温的响应

以青藏高原高寒草甸和高寒沼泽中的两种优势物种小嵩草(Kobresia pygmaea)和藏嵩草(Kobresia tibetica)为研究对象,采用开顶式增温室(OTCs)模拟气候变暖,对比分析两种植物叶片形态和解剖结构特征、根活性及地上—地下部分化学计量特征对增温的响应差异。结果表明:增温显著增加了小嵩草叶片的长度和叶片的数量,也显著增加了藏嵩草株高和叶片长度;增温没有明显改变小嵩草和藏嵩草的叶片上表皮厚度、下表皮厚度、下表皮细胞角质层厚度、叶肉细胞长和叶肉细胞宽;增温增加了小嵩草根系活跃吸收面积,对小嵩草和藏嵩草其他根系活性指标没有显著影响;增温降低了小嵩草地上部分N含量,对小嵩草地上部分C、P含量没有影响;增温降低了藏嵩草地上部分C、N含量,对P含量没有影响;增温增加了小嵩草和藏嵩草地上部分C/N比,提高了两种优势植物对氮素的长期利用效率;增温对小嵩草地下部分化学计量学特征没有影响,而降低了藏嵩草地下部分C含量和C/N比。     

QTL mapping reveals a two-step model for the evolutionary reduction of inner microsporangia within the asteracean genus <Emphasis Type="Italic">Microseris</Emphasis>

The reduction of inner (adaxial) pollen sacs (microsporangia, MS) as a diagnostic character for the three asteracean species, Microseris bigelovii, Microseris elegans and Microseris pygmaea, was analysed in an interspecific cross between Microseris douglasii and Microseris bigelovii with 4 MS and 2 MS, respectively, using the average number of MS per plant as a quantitative character. A previous QTL (Quantitative Trait Locus) analysis had revealed one major QTL (3B) and three modifier QTLs (3A, 4A, 7A) with epistatic effects only on the homozygous recessive 2 MS genotype of QTL 3B. Here we performed a bulked segregant analysis on four 2 MS and four 4 MS DNA-bulks with 407 EcoRI/MseI AFLP-primer combinations each. In this way additional AFLP markers were mapped close to QTL 3B and QTL 3A. Three of them were converted to SCAR (Sequence Characterized Amplified region) markers. All markers were tested in natural populations of the disporangiate (2 MS) species M. bigelovii, M. elegans and M. pygmaea, and in different populations of tetrasporangiate (4 MS) M. douglasii. The marker distribution suggests that locus 3B mutated in a progenitor of the disporangiate species. QTL 3A has evolved in the 2 MS background of the major gene in the disporangiate species. Since M. pygmaea and M. bigelovii are the sister group to M. elegans, the 4 MS genotype for (markers of) QTL 3A in M. pygmaea populations is most likely due to a back mutation to the 4 MS state and could explain the slight instability of the 2 MS phenotype in this species.Communicated by O. Savolainen     

Nuclear DNA markers of the Australian tetraploid Microseris scapigera and its North American diploid relatives

The allotetraploid lactucean Microseris scapigera of Australia and New Zealand has presumably arisen in western North America by hybridization between an annual and a perennial diploid species followed by polyploidization and long-distance dispersal. A phylogenetic tree of various North American diploids, based on RFLPs in the nuclear DNA, confirmed the division of the genus into a clade containing the diploid annuals and a clade containing the diploid perennials. Four RFLP markers were shared among all accessions of M. scapigera and all the diploid accessions. Twelve markers found in the outgroup (Uropappus lindleyi) were absent in all Microseris. A cladogram of plants from six populations of M. scapigera based on eight RFLP markers shows a progressive specialization of three clades of two populations each. Two populations without any markers differentiating them from the North American diploids form the basic clade. These consist of plants with an apparently derived morphology that are self-compatible (or agamospermic) and thereby differ from most M. scapigera. Few markers in M. scapigera could be attributed to one or the other parental genome. As yet, we have found only one ITS 1 sequence of the nuclear ribosomal cistrons in M. scapigera. This sequence has features of both parental sequences.     

Modifiers of heterocarpy determine capitulum size inMicroseris hybrid D 14 (Asteraceae — Lactuceae)

The proportion of the outer, “hairy” achenes in capitula of hybrid D 14 (Microseris pygmaea × M. bigelovii) is determined by the same major gene as in its sister hybrid, B 87, as shown by marker segregation. Crossover between major gene and markers shows their genetic independence. Two modifiers segregating 9:7 influence the proportion of hairies in plants homozygous for the major gene. These same modifiers (or two genes linked with them) also determine the segregation of the number of achenes per head. Most likely, the modifiers act indirectly via a residual dependence of heterocarpy on capitulum size within strict quantitative limits set by the major gene. The identification of modifiers in a polygenic system as major genes for another character acting pleiotropically supports our contention that relatively few genes interact in plant development to determine key morphological characteristics at the organismic level.     

The karyotypes ofMicroseris bigelovii,M. douglasii,andM. pygmaea (Asteraceae)

The karyotypes of the three annuals,Microseris bigelovii, M. douglasii andM. pygmaea, consist of 2n = 18, small, submetacentric chromosomes. Length, centromere position, C-banding pattern, silver staining of NOR's, and the use of base specific fluorochromes, allow the identification of four of the nine chromosome pairs. The banding pattern ofM. bigelovii andM. pygmaea is identical, but intraspecific differences are found between strains ofM. douglasii.     

Ecology of achene dimorphism in Leontodon saxatilis

BACKGROUND [corrected] AND AIMS: Leontodon saxatilis produces two morphologically distinct achenes (morphs) in a single capitulum: one row of dark brown achenes without a pappus lies at the edge ('peripheral achenes'; 0.74 +/- 0.18 mg) while the inner ones are light brown with a pappus ('central achenes'; 0.38 +/- 0.07 mg). The hypothesis that achene heteromorphism in L. saxatilis widens its ecological amplitude was tested. KEY RESULTS: Achenes of both morphs germinated over the same range of temperatures (6-33 degrees C) but the central achenes showed significantly higher germination percentages, and the two also differed significantly in their annual dormancy cycle, with the peripheral achenes showing greater dormancy for part of the year. Seedlings from the two morphs did not differ significantly in total biomass after 2 and 4 weeks of growth, neither did they differ significantly in root and shoot weight and root:shoot ratio. Plants from both morphs growing at different regimes of soil moisture, nutrients and competition did not differ significantly in their number of achenes per capitulum. While the number of central achenes varied, that of peripheral achenes remained constant at approx. 13. Drier soil led to an increase in the number of central achenes in plants from both morphs. CONCLUSIONS: The peripheral achenes can replace the mother plant in the following growing season, whereas the central achenes are well adapted for wind dispersal and thus for colonization of new sites. However, the extent of variation in germination characteristics was similar to that found in seed populations of homomorphic plants, which suggests that germination percentage and other growth characteristics do not contribute to widening the ecological amplitude. The habitat of most heteromorphic species, the morphs of which differ greatly in germination and/or growth characteristic, are deserts or highly disturbed areas where such differences are highly advantageous, but in the moderate habitat of L. saxatilis the differences may prove a disadvantage.     

重庆荨麻科一新种——缙云赤车

在重庆市北碚区缙云山野外植物调查过程中,于海拔500～800 m的溪边阴湿沟谷中发现一种荨麻科植物,虽然该物种雌花序具花序托,但因其雌花花被片5枚,外面先端具角状突起,瘦果只具瘤状突起,更符合赤车属的特征,且结合分子系统学证据,该物种明显属于赤车属.该物种的特征在于其具花序托和总苞的雌花序,根据这一特征可将其置于头序赤...