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.     

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.     

Phenotypic Plasticity of Capitulum Morphogenesis in Microseris pygmaea (Asteraceae: Lactuceae)

Inbred populations of the annual Chilean species Microserispygmaea show phenotypic plasticity in the number of floretsper capitulum. In order to find out how this plastic variationmay arise during development, two inbred lines (A92 and C96)were grown under short day conditions. Groups of plants fromeach strain were transferred to long day conditions at about2 week intervals. In this way we introduced variation for plantsize (number of leaves per plant) at onset of flowering. Thenumber of florets per mature capitulum increased linearly withplant size. After transfer to long day conditions, plants wereharvested daily for light microscopic measurements of meristemwhole mounts. Only the first capitulum of each plant was analysed.All florets were formed after 12 d in strain C96 and after 14d in strain A92. In order to detect the effect of plant sizeon morphogenesis, we performed a multiple regression analysisof developmental parameters on time and number of leaves. Widthand height of the capitulum receptacle increased daily witha growth rate depending on the starting size. Differences inmeristem size were detected already in vegetative plants ofdifferent sizes. In contrast, there was no influence of plantsize on floret primordium size. We combined the multiple regressionmodels in one simple model for prediction of floret numbersfrom numbers of leaves per plant at onset of flowering. Predictionsof this model agree with observed relationships in both inbredlines.Copyright 1994, 1999 Academic Press Asteraceae, capitulum, inflorescence, meristem, meristic character, Microseris pygmaea, morphogenesis, phenotypic plasticity     

Pappus part number in annual species ofMicroseris (Compositae,Cichoriaceae)

All North American annual species of the genusMicroseris have a five-part pappus, the one South American annual,M. pygmaea, has ten pappus parts. The pappus develops over a constant number of ten provascular bundles with or without inhibition between alternate sites of pappus development. Each natural population contains a predictable proportion of achenes with aberrant pappus part numbers. Hybridization betweenM. bigelovii (5 parts) andM. pygmaea results in F 1 and F 2 plants with many aberrant achenes. In each plant either five or ten can be shown to be the basic number with aberrant numbers following a Poisson distribution for numbers added to 5 or deleted from 10. Occasional plants show no basic number but have a random distribution of numbers about an intermediate mean. The evolutionary genetics of this character is discussed.     

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.     

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.     

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.     

Comparison of chloroplast and nuclear phylogeny in the autogamous annualMicroseris douglasii (Asteraceae: Lactuceae)

Morphology suggests that the Californian annualMicroseris douglasii is a monophyletic sister group to the other three diploid annuals ofMicroseris. Phylogenetic analysis of 44 inbred strains ofM. douglasii derived from 23 populations with 72 RAPD markers in the nuclear DNA strongly supports this phylogeny. However, 13 chloroplast RFLPs divideM. douglasii into four distinct groups. Two of these each share one or more cpRFLPs withM. bigelovii andM. pygmaea. Several hypotheses can explain the incongruence between nuclear and chloroplast phylogeny: (1) random sorting out of chloroplasts during phylogeny from a polymorphic pool, (2) cytoplasmic introgression from the related annualM. bigelovii intoM. douglasii after hybridization followed by elimination of theM. bigelovii nuclear genome. We suggest cytoplasmic introgression as the most likely origin. Possible remnants of nuclear introgression have been found in two populations ofM. douglasii that are polymorphic for chloroplast types. In these populationsM. bigelovii type chloroplast DNA seems to be accompanied by nuclear genes for flower color and leaf shape.     

Genetic analysis of aMicroseris douglasii (Asteraceae) population polymorphic for an alien chloroplast type

Recent evidence suggests chloroplast introgression fromMicroseris bigelovii intoM. douglasii. We have examined 23 plants from a population ofM. douglasii polymorphic forM. douglasii andM. bigelovii chloroplast types. All 23 plants were completely homozygous for morphological and RAPD markers, and inbred lines derived by selfing have been used for DNA analysis. Chloroplast RFLP analysis identified 16 plants withM. bigelovii chloroplasts and seven withM. douglasii chloroplasts. The nuclear genomes of the 16 plants withM. bigelovii chloroplasts were examined with 22 primers for RAPD amplification products shared exclusively withM. bigelovii. Five of 268 markers appeared to be shared betweenM. bigelovii and one or more of these 16 plants on the basis of their position in gels. Detailed examination of these five amplification products showed that none of them are nuclear DNA fromM. bigelovii. Very little, if any, nuclear DNA fromM. bigelovii can be present inM. douglasii plants with chloroplasts typical ofM. bigelovii. The study demonstrates the usefulness of the RAPD technique for screening large numbers of markers to select a few potentially informative ones for rigorous examination.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

The Effect of Rooting Space on Whole Plant and Leaf Growth in Brussels Sprouts (Brassica oleracea var. gemmifera)

Seedlings of Brassica oleracea var. gemmifera DC. (Brusselssprouts) were grown in four pot sizes over a 4-week period.Whole plant, stem, root and foliage d. wts and foliage area,together with specific leaf area, leaf area ratio and numberof leaves initiated were reduced by restricting rooting space.Individual leaves showed similar reductions in d. wt and area,with the effect being more pronounced in later-formed leaves.Cell counts and measurements on the epidermis and palisade mesophylllayers of the first four leaves showed that the reduction ingrowth was due to reduced cell division. Cell numbers in thefirst-formed leaf were halved over the range of pot sizes used,and there was a progressively greater reduction in cell numbersin later-formed leaves. There was some tendency for cell sizeto decrease with decreasing rooting space, but this was notgeneral and was most marked between plants grown in the twosmallest pot sizes. Brassica oleracea var. gemmifera, Brussels sprouts, rooting space, growth analysis, leaf growth, cell numbers, cell sizes     

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     

Comparisons of Earlier- and Later-formed Pods of Chickpeas (Cicer arietinum L.)

In chickpeas (Cicer arietinum L.) flowering and pod developmentproceed acropetally. In plants grown under normal field conditionsat Hyderabad, in peninsular India, and at Hissar in north India,at successively apical nodes of the branches there was a declinein pod number per node, weight per pod, seed number per podand/or weight per seed. The percentage of nitrogen in the seedswas the same in earlier and later-formed pods at Hyderabad;at Hissar the later-formed seeds contained a higher percentage.Earlier- and later-formed flowers contained similar numbersof ovules. The decline in seed number and/or weight per seedin the later-formed pods of 28 out of 29 cultivars indicatedthat pod-filling was limited by the supply of assimilates orother nutrients. By contrast, in one exceptionally small-seededcultivar there was no decline in the number or weight of seedsin later-formed pods, indicating that yield was limited by ‘sink’size. Cicer arietinum L., chickpea, flowering, pod development, seed number, seed weight, nitrogen content     

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.     

Evolution of microsporangium numbers in Microseris (Asteraceae: Lactuceae)

The genus Microseris contains species with disporangiate stamens and species with tetrasporangiate stamens. We determined the number of microsporangia per stamen in serial sections of heads from all 13 species of Microseris, its close relative Uropappus lindleyi, and the two allopolyploid species of Slebbinsoseris. Four Microseris species, three diploid and one tetraploid, have two microsporangia per stamen; all other species investigated have four. The most parsimonious assumption is that the disporangiate condition is derived and arose once in the evolution of Microseris. The inheritance of the number of microsporangia per stamen in crosses between M. bigelovii (disporangiate) and M. douglasii (tetrasporangiate) was determined. Segregation of microsporangium number per stamen in F2s derived from these crosses is quantitative rather than Mendelian. The average number of microsporangia per stamen in the F2 plants ranges from 2.0 to 4.0. There is a predominance of tetrasporangiate stamens in the F1 and in most F2 plants. The observed pattern of inheritance suggests a major gene with dominance and quantitative modifiers.     

Chloroplast and nuclear DNA variation among homozygous plants in a population of the autogamous annualMicroseris douglasii (Asteraceae,Lactuceae)

The autogamous diploid annualMicroseris douglasii of California occurs in many isolated populations. The populations consist of one to many highly inbred biotypes. Morphological variation among populations usually is greater than within populations. In spite of the virtual absence of gene flow even within populations, genetically determined character differences are randomly distributed and associated throughout the range of the species. Recent evidence even suggests introgression of chloroplasts from the relatedM. bigelovii. Offspring families from 25 plants of a very variable population were raised and examined for segregation of morphological and molecular (RAPD) markers. All 25 original plants were completely homozygous for all markers, but each differed from all others at least in some markers. The population consisted of two genetically isolated groups of plants: a distinct inbred line (3 plants) and 22 plants with random associations of a common set of markers and characters, possibly recombinant inbreds from a past hybridization event. One of these 22 plants contained a chloroplast genome found inM. bigelovii, the other 24 plants a chloroplast genome found only inM. douglasii.     

Comparative Response of Nodulated and Nitrogen-Supplied Cowpea (Vigna unguiculata (L.) Walp. var. Tuy) Plants to Infection by Cowpea Mosaic Virus

The effect of infection by the Cowpea Mosaic Virus (CpMV) onseveral parameters relevant to symbiotic nitrogen fixation wasdetermined in cowpea (Vigna unguiculata (L.) Walp. var. Tuy)plants nodulated with two strains of Rhizobium cowpea: IVIC–124and IVIC–38. Plants were virus-infected at the seedlingstage before Rhizobium inoculation. The effect of CpMV infectionon plant growth was analysed in nodulated and nitrogen-suppliedplants at 18, 25 and 35 d after germination. At all developmentalstages of nodulated plants CpMV infection caused a reductionof leaf chlorophyll content, leaf area, dry weight of shootsand roots, total nodule weight and nodule number. Most of thenodules from 18- and 25-d-old CpMV-infected plants did not exhibitleghaemoglobin pigmentation. CpMV infection delayed the onsetof nitrogenase activity in nodules of the rhizobial strain IVIC–124and the enzyme activity measured on a per plant basis was reducedin both strains at the first and second harvests. Significantnitrogenase activity was detected in 35-d-old infected plants.Some of the nodules of the rhizobial strain IVIC-124 and mostof the nodules from plants nodulated with the strain IVIC-38developed leghaemoglobin; however, the nodule-specific nitrogenaseactivity, estimated on a milligram nodule dry weight basis,was always higher in virus-infected plants, particularly in18-d-old CpMV-infected plants harbouring the IVIC–124strain. CpMV-infected nodules had a larger peribacteroidal space,a reduced number of peribacteroid units, a greater number ofbacteroids per unit, a lower number of vesicles and 88% lowertotal reducing sugar content. Starch accumulation was detectedin infected leaves of nodulated plants during the first harvest,while high levels of leaf reducing sugars and protein were presentat the second harvest. In healthy nodulated plants the rhizobialstrain IVIC–124 was shown to be more efficient than IVIC–38in promoting plant growth. However, the results indicate thatnodulation by rhizobial strain IVIC–124 and growth ofplants harbouring this strain were affected to a greater extentby virus infection. The effect of CpMV infection on leaf chlorophyllcontent, leaf area, carbohydrate level, leaf proteins and growthof nitrogen-supplied plants, as well as the symptoms inducedin the leaves, were less conspicuous than in nodulated plants. Key words: Cowpea, Rhizobium, virus infection, nodule untrastructure     

Homeotic, Meristic and Cytological Floral Mutants of Thryptomene calycina (family Myrtaceae)

Twenty plants with various phenotypic abnormalities to the flowerswere selected from very large populations of Thryptomene calycinain the Grampian and Black Ranges. Most of these had impairedreproductive function. Normal flowers were epigynous with fivesepals, five petals, five anthers, a single style and two anatropousovules. The mutants were two partially male sterile, tetraploidplants with large flowers, one of which occasionally producedadditional flowers from the leaf axils with peduncles as wellas pedicels; one plant which produced a proportion of hexapetaloidflowers with six stamens; three gross mutants with fleshy, bracteoidpointed petals and sepals, no stamens, vestigial styles andstigmas, exposed ovules and no inferior ovary; one plant withfleshly, bracteoid pointed sepals, vestigial style and stigmabut with exposed ovular structures replaced by four to fivesterile ovules generally inside an abnormal ovary; two plantswith reduced ovary diameter and sterile ovules, shortened style,five reduced sepals and petals and five to eight anthers; threeanthocyanin-free plants; three plants with pink sepals; twoplants with half-sized flowers which produced a proportion offasciated stems; one plant which occasionally produced flowerswithout pedicels which virtually resulted in organs which wereleaf-flower composites; two plants which produced sepals andpetals which contained chlorophyll and prematurely senesced,and had partial substitution of petals by anthers.Copyright1993, 1999 Academic Press Thryptomene calycina, Myrtaceae, Victorian lace flower, floral mutations, mutants, homeotic, meristic, tetraploid, fasciation, male sterility, cut flowers     

The Evolutionary Reduction of Microsporangia in Microseris (Asteraceae): Transition Genotypes and Phenotypes

Abstract: The loss of the two inner (adaxial) microsporangia (MS) on the anthers is a shared, derived character for three species of the genus Microseris (Asteraceae). In a hybrid between M. douglasii (4 MS) and M. bigelovii (2 MS), one major gene and four modifier loci are responsible for the difference in MS number. The homozygous recessive (2 MS) genotype of the major gene is necessary but not sufficient for the reduction. In addition, at least five M. bigelovii (2 MS) alleles of the three major modifiers are needed for a stable 2‐MS phenotype in all florets of a plant. One, two or three M. bigelovii alleles of the modifiers cause the random reduction or loss of some of the adaxial MS. When the major gene and two modifiers specify 2 MS and only one modifier is homozygous for the M. douglasii (4 MS) alleles, sister plants can have any phenotype from pure 2 MS to pure 4 MS. Here, we examine the phenotypic expression of these genotypes raised under the normal winter annual conditions and under long‐day conditions. In all cases, the phenotypes vary among sister plants, but the range of variation (most notably under long‐day conditions) depends on the specific modifier gene contributing the M. douglasii alleles. The phenotypic variance in one of the genotypes was decreased by a factor of ten in the depauperate heads produced in the long‐day experiment. This effect is mediated by a dependence of the MS phenotype on the position of the floret relative to the edge of the flowering head (capitulum) and directly by the size of the capitulum. Genotypes specifying phenotypes with more or less precisely two or four MS in all florets show hardly any dependence on environmental or developmental factors. The significance of these observations lies in the non‐linear, “canalized” relationship between phenotypic expression and gene dosage, which shows how a qualitative morphological change dependent on a single major gene mutation can pass through a potentially maladaptive intermediate stage.     

The Assessment of Speckled Leaf Blotch in Winter Wheat in New Zealand

Disease symptoms of speckled leaf blotch (Mycosphaerella graminicola)were first observed in early August, 86 d after sowing a winterwheat field trial in New Zealand. Disease hastened senescenceof all leaves, delayed the expansion of new leaves during springand reduced the maximum size of some later-formed leaves. Theabsolute green leaf area was reduced by disease through to fullsenescence of leaves, which occurred one week earlier than inhealthy plants. Disease indices calculated from the data onpercentage diseased area alone indicated a decline in diseaseduring rapid growth of the plants in spring as the temperatureincreased. However, this apparent decline was generated by themethod of calculation and was not evident when effects on leafarea were also considered. Senescence induced by the pathogenwas an important aspect of the disease syndrome. Mycosphaerella graminicola, Septoria tritici, speckled leaf blotch, winter wheat, yield loss     

取食不同寄主植物对棉蚜后代抗药性的影响

测定了5种药剂对棉蚜Aphis gossypii抗氰戊菊酯、吡虫啉品系和敏感品系取食棉花、黄瓜和石榴的后代的毒力,并对它们的后代体内乙酰胆碱酯酶和羧酸酯酶的比活力做了初步探索。结果表明,氰戊菊酯抗性品系取食棉花比取食黄瓜的后代对氰戊菊酯的抗性大76.4倍,对灭多威、氧乐果、硫丹和吡虫啉的抗性也大0.5～4.6倍;取食石榴的后代对5种药剂的抗性介于取食棉花和黄瓜的之间。吡虫啉抗性品系的测定结果与氰戊菊酯抗性品系基本一致。敏感品系取食黄瓜比取食棉花的后代对5种药剂的敏感性更高。3个品系取食不同植物的后代相比,其体内乙酰胆碱酯酶的比活力,取食棉花的为取食黄瓜的2.4～2.8倍;羧酸酯酶的比活力,取食棉花的为取食黄瓜的1.8～2.4倍。证明棉蚜的抗性和敏感品系取食的寄主植物不同,可引起对药剂敏感性的变化。乙酰胆碱酯酶和羧酸酯酶活力的变化均是引起这种变化的重要因素。