The effect of resistant solanaceous plants on the sex ratio of Heterodera rostochiensis and the use of the sex ratio to assess the frequency and genetic constitution of pathotypes

The sex ratios of two Heterodera rostochiensis populations were compared on fifteen solanaceous plants. Larvae did not mature in the roots of Atropa belladona, Datura stramonium or Nicotiana tabacum but adults were produced on the other plants, on some only males and on others, males and females. With two other populations, but not with a third, the percentage of larvae that became female on a potato hybrid with a dominant gene for resistance derived from Solanum tuberosum ssp. andigena increased during successive generations on the hybrid in accordance with the hypothesis that such larvae are double recessives (aa). Fewer larvae became adult in plants with two dominant genes for resistance than in plants with one, and such plants probably exerted selection of both males and females. A population from St Brelades (Jersey) produced fewer females and more males on the susceptible variety Arran Banner than did either the Thorney (Cambs.) or Aberystwyth populations. A much greater percentage of larvae from the St Brelades population became female on an ex andigena hybrid derived from CPC 1690 than on one derived from CPC 1685.     

The population genetics of the potato cyst-nematode, Heterodera rostochiensis: mathematical models to simulate the effects of growing eelworm -resistant potatoes bred from Solatium tuberosum ssp. andigena

To follow population changes when potato varieties with resistance to Heterodera rostochiensis derived from Solanum tuberosum ssp. andigena were grown on infested land, a computer programme was written including three mathematical relationships: (1) a law relating multiplication to pre-cropping density; (2) two mathematical models of inheritance of ability to overcome resistance; (3) a law relating the proportion of larvae able to become female to pre-cropping population density. The programme also included four parameters: (1) the maximum possible reproductive rate; (2) the fraction of the population (eggs) not participating in reproduction when potatoes are grown and carried over to the following year unchanged; (3) the fraction carried over annually when other crops are grown; (4) the frequency of larvae able to become female in the population initially. Population density was measured relative to the equilibrium density and was therefore independent of the units in which density is usually measured. After supplying a range of parameters for all the above to include those likely to be encountered in practice, the changes expected (a) in the frequency of larvae able to become female in the roots of resistant varieties and (b) in population density were computed for resistant varieties grown continuously or alternately with susceptible varieties in crop rotations of different lengths. Because well established field populations are relatively dense, observed reproductive rates are small and rarely approach the maximum possible. Reproductive rate is therefore a relatively unimportant determinant of genetic change. The fraction of the population carried over to the following year is more important because it affects the length of a crop rotation necessary to make loss of potato yield acceptable, determines what the multiplication rate will be and influences the speed of genetic change by providing a reservoir of initial type males which backcross with any genetically different females that may develop on the roots of resistant plants. No experiment seems to have been done specifically to determine the parameters needed to calculate population changes. Some values can be obtained from the literature but mostly they must be guessed. When the law relating the proportion of larvae able to become female to pre-cropping population density was included in the computations, it had little effect initially but later, after several generations, it delayed genetic change. Two field experiments, one by Huijsman (1961) another by Williams (1958) and Cole & Howard (1962 a), provide some of the variables needed to compute trends in population density. Best fitting variables were computed for the data in these experiments by the method of maximum likelihood. The computed parameters for one experiment were not very realistic but those for the other were in line with what would be expected in practice and tended to favour the hypothesis that larvae able to become female in the roots of resistant plants are double recessives (aa). The computations lead us to suggest that the best policy for potato growers who have fields suitable for resistant varieties is to alternate resistant with susceptible varieties in a crop rotation containing potatoes every 3 or 4 years.     

Disk electrophoresis of proteins of Heterodera species and pathotypes of Heterodera rostochiensis

The proteins extracted by aqueous buffer from the females of six Heterodera spp. were separated into bands by polyacrylamide gel electrophoresis and the patterns compared. Heterodera schachtii and H. trifolii, with lemonshaped cysts, had a similar pattern of bands, differing greatly from that of the round-cyst nematodes H. virginiae and H. rostochiensis. Ten pathotype A populations of H. rostochiensis had similar patterns, distinguishable from those of twelve pathotype E populations by the absence of two bands and by the presence of another rarely found in pathotype E. Bands from pathotype B populations were indistinguishable from those of pathotype E. No consistent difference was detected between populations of the same pathotype. These results support the suggestion by Jones, Carpenter, Parrott, Stone & Trudgill (1970) that H. rostochiensis Woll. comprises two species.     

Breeding for resistance to the white potato cyst-nematode, Heterodera pallida

Progenies bred from material derived from the wild potato, Solanum vernei and from the cultivated potato, S. tuberosum ssp. andigena, clone CPC 2775, were compared for their resistance to Heterodera pallida, pathotype E. The influence of additional resistance derived from the wild species, S. multidissectum, was also investigated. Both S. vernei and CPC 2775 gave progenies with variable levels of resistance and there was often no clear segregation into resistant and susceptible categories. Incorporation of gene H₂ derived from S. multidissectum increased resistance to pathotype E of H. pallida for resistant material bred from both S. vernei and clone CPC 2775. The results indicate that adequate resistance to all British populations of potato cyst-nematodes can best be obtained by combining the factors for resistance from the two Andigena clones, CPC 1673 (gene H₁) and CPC 2775 (gene H₃), and from S. multidissectum (gene H₂).     

The pathogenicity and increase of Heterodera rostochiensis on tomato cultivars, self-rooted or grafted on to rootstocks.

Self-rooted tomatoes ‘Ailsa Craig’ and ‘Kingley Cross’, and scions of ‘Ailsa Craig’ grafted on to different rootstocks, were grown in soil infested with different numbers of Heterodera rostochiensis (Mainly pathotype A). Three rootstocks (RS₄, RSKN, ISKVF) tolerated damage from nematode populations of up to 125 eggs/g soil, but nematodes increased up to x45 on them. Two nematode-resistant F₁ hybrid rootstocks, B66₃₃ (ex Lycopersicon pimpinellifolium B6₁₇₃) and B6₀₁₅ (ex L. hirsutum var. glabratum B6₀₁₃), tolerated up to 125 eggs/g soil; nematode populations declined as much as 80% under B66₃₃, and increased little under B6₀₁₅. On the self-rooted ‘Ailsa Craig’ and ‘Kingley Cross' the numbers of nematodes increased markedly (maximum increase x40), and greatly affected growth and yield so that initial populations greater than 62 eggs/g soil caused crop failure.     

Microcosm studies on intraguild predation between female erigonid spiders and lacewing larvae and influence of single versus multiple predators on cereal aphids

Abstract: Intraguild predation between female erigonid spiders [Erigone atra (Blackwall) and Oedothorax apicatus (Blackwall), Araneae, Erigonidae] and lacewing larvae (second instar larvae of Chrysoperla carnea (Stephens), Neuropt., Chrysopidae) and interaction effects of predator combinations on cereal aphids were investigated in a microcosm system under laboratory conditions. The microcosm experiments were run for 7 days and consisted of 15wheat seedlings, 15 Sitobion avenae (F) (Hom., Aphididae) as start population, plus a female spider or a lacewing larva or a combination of a spider plus a lacewing larva. The mortality rate of lacewing larvae was significantly increased by 44 and 31% due to intraguild predation by female spiders of E. atra and O. apicatus in comparison with lacewing larvae that were kept alone. The final aphid numbers in the microcosms were significantly reduced by all single predator treatments (spiders, lacewing larvae) and the predator combinations in comparison with controls without predators. The predation effect on aphid populations due to both spider species was similar and not statistically different. An additive effect of the predator combinations ‘spider plus surviving lacewing larva’ was found for both spider species resulting in reduced aphid numbers compared with the single predator treatments. When the lacewing larva was killed by an E. atra female the effects on aphids were non‐additive, but aphid numbers were not statistically increased compared with the lacewing larva treatment. When the lacewing larva was killed by an O. apicatus female, the effects of spider and C. carnea larva were additive on aphid numbers. In the presence of additional prey (fruit flies and Collembola) intraguild predation was not found and E. atra females had no significant effect on the survival of lacewing larvae. In addition, E. atra females had no significant effect on aphid numbers in the presence of fruit flies and Collembola, but in combination with a lacewing larva that survived, a significantly greater reduction of the aphid population was observed compared with the lacewing larva treatment. The body mass of lacewing larvae at the end of the experiment was not statistically influenced by the presence or absence of an E. atra female.     

Hybridization studies of Haemonchus contortus (rudolphi, 1803) and H. placei (place, 1893) (Nematoda: Trichostrongylidae)

Hybridization experiments between Haemonchus contortus from sheep and H. placei from cattle indicate that the H. placei vulvar morph type and the inability of the eggs to hatch and develop at 11°C were inherited as dominant traits in the hybrids. The size of the third stage larvae was similar to H. placei in hybrids from the mating of male H. placei × female H. contortus while larvae from the reciprocal mating were intermediate in size. Hybrids produced by the mating of thiabendazole resistant female H. contortus × non-resistant male H. placei were also resistant. The F₁ males of the mating between male H. contortus and female H. placei were sterile. Male sterility did not occur in the reciprocal cross until the F₂ generation. Female hybrids from these generations had a low level of fertility when backcrossed to males of either parent species. Cytological studies of the hybrid males indicated that sterility was due to several kinds of meiotic disturbance and that spermatogenesis stopped during metaphase I. The chromosomes in eggs of unfertilized females did not undergo meiosis and polar body formation; instead they increased in number by a process of endomitosis. Unfertilized eggs in the faeces were characterized by uneven cytoplasmic division and abnormal shape. It is proposed that hybrid sterility can lead to the local eradication of one species of Haemonchus by the other. Furthermore, the outcome of which species is eradicated can be influenced by such variables as initial population sizes, climate, and the local ratio of sheep to cattle.     

REPRODUCTIVE ISOLATION IN RHABDITIDAE (NEMATODA: SECERNENTEA); MECHANISMS THAT ISOLATE SIX SPECIES OF THREE GENERA

We have attempted interspecific hybridizations among six species of rhabditid nematodes: Caenorhabditis elegans, Caenorhabditis briggsae, Caenorhabditis remanei, Caenorhabditis sp. v, Rhabditis sp., and Pelodera teres. Copulation was observed in all crosses between Caenorhabditis species; however, none resulted in the generation of stable hybrid populations. No copulation was observed in crosses between Caenorhabditis males and Rhabditis or Pelodera females, even when congeneric females were present, suggesting that Caenorhabditis males are able to selectively recognize congeneric females by a short-range stimulus. All pairwise combinations of Caenorhabditis species were isolated to some degree by gametic mechanisms; 7 of 12 combinations were cross infertile and 5 of 12 were cross-fertile but had low brood sizes. In cross-fertile combinations, most hybrid embryos were inviable and arrested prior to gastrulation. Only in crosses of C. briggsae males to C. sp. v females did any hybrids survive embryogenesis. Most of these C. briggsae/C. sp. v hybrids arrested during larval development, and the few that reached adulthood invariably were female. These results are consistent with the presence of at least two lethal factors in the C. briggsae-C. sp. v combination: a maternal lethal factor in the cytoplasm of C. briggsae and a recessive lethal factor on the X chromosome of C. sp. v.     

Selection for virulence in the potato cyst-nematode, Globodera pallida

The ability of six populations of Globodera pallida to multiply on Solanum vernei hybrids (Morag, 12380 2, Glenna and Santé) and S. tuberosum andigena hybrids (A 27/23, Paladin (ZB 35/29) and ZC 83/6) was studied in microplots and pots. Evidence was obtained of selection for virulence in one population after one year's cultivation of cvs Glenna or Morag. Similarly, after four years' continuous cultivation of cvs Morag or ZC 83/6 in microplots, three of five populations of G. pallida contained more virulent nematodes than the same populations maintained for the same period on cv. Pentland Crown (susceptible to G. pallida). These results are discussed in relation to earlier work and the use of partially resistant cultivars in the integrated control of G. pallida.     

Influence of humidity and a surfactant-polymer-formulation on the control potential of the entomopathogenic nematode <Emphasis Type="Italic">Steinernema feltiae</Emphasis> against diapausing codling moth larvae (<Emphasis Type="Italic">Cydia pomonella</Emphasis> L.) (Lepidoptera: Tortricidae)

The codling moth (Cydia pomonella L.) is a serious pest of pome fruit. Diapausing cocooned larvae overwinter in cryptic habitats in the soil or in the bark of infested trees. The entomopathogenic nematode Steinernema feltiae (Filipjev) (Rhabditida: Steinernematidae) is used to control diapausing codling moth larvae. The objective of this study was to define environmental conditions favouring the performance of the nematodes. Cocooned larvae were more susceptible than non-cocooned larvae. Susceptibility of pupae was low. To determine the influence of decreasing water activity (a_w-value) on the activity of the nematodes, mortality of codling moth larvae and Galleria mellonella L. were tested in sand-sodium-polyacrylate mixtures of variable water activity. S. feltiae was able to infect both insects at a_w-values >0.9. Cocooned larvae of both insects died at lower a_w-values than non-cocooned larvae. Mortality of cocooned larvae did not further increase after half an hour of exposure to nematodes, whereas the mortality of non-cocooned larvae increased with increasing exposure time. LC₅₀ and LC₉₀ considerably decrease with increasing RH. The negative influence of the relative humidity (macro environment) was less important than the effect of the water activity in the bark substrate (micro environment). The micro environment can be manipulated by applying S. feltiae with higher volumes of water. A surfactant-polymer-formulation significantly increased nematode efficacy and can buffer detrimental environmental effects.     

Measuring the discrepancy between fecundity and lifetime reproductive success in a pollinating fig wasp

Lifetime reproductive success in female insects is often egg‐ or time‐limited. For instance in pro‐ovigenic species, when oviposition sites are abundant, females may quickly become devoid of eggs. Conversely, in the absence of suitable oviposition sites, females may die before laying all of their eggs. In pollinating fig wasps (Hymenoptera: Agaonidae), each species has an obligate mutualism with its host fig tree species [Ficus spp. (Moraceae)]. These pro‐ovigenic wasps oviposit in individual ovaries within the inflorescences of monoecious Ficus (syconia, or ‘figs’), which contain many flowers. Each female flower can thus become a seed or be converted into a wasp gall. The mystery is that the wasps never oviposit in all fig ovaries, even when a fig contains enough wasp females with enough eggs to do so. The failure of all wasps to translate all of their eggs into offspring clearly contributes to mutualism persistence, but the underlying causal mechanisms are unclear. We found in an undescribed Brazilian Pegoscapus wasp population that the lifetime reproductive success of lone foundresses was relatively unaffected by constraints on oviposition. The number of offspring produced by lone foundresses experimentally introduced into receptive figs was generally lower than the numbers of eggs carried, despite the fact that the wasps were able to lay all or most of their eggs. Because we excluded any effects of intraspecific competitors and parasitic non‐pollinating wasps, our data suggest that some pollinators produce few offspring because some of their eggs or larvae are unviable or are victims of plant defences.     

Eucalypt hybrid susceptibility to Gonipterus scutellatus (Coleoptera: Curculionidae)

Abstract Damage caused by the eucalypt snout weevil Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae) (=G.gibberus Boisduval) was found on a greater proportion of F₁Eucalyptus amygdalina Labill. X E. risdonii J. D. Hook, hybrid trees than either of the pure species, in a replicated field trial in south‐east Tasmania. A greater proportion of E. risdonii trees was also damaged than E. amygdalina trees. A study of the pattern of oviposition within the trial revealed no difference in oviposition by G. scutellatus between E. risdonii and E. amygdalina. Oviposition by G. scutellatus was significantly higher on the F₁ and F₂interspecific hybrids between these two eucalypt species compared with pure species crosses. There were no apparent differences in damage or oviposition levels between the F₁ hybrids and the single large F₂ progeny included in the trial. This finding provides little evidence that hybrid susceptibility is due to hybrid breakdown after the first generation. Rather, the evidence is consistent with the hypothesis that there may be different mechanisms of host defence operating in each species that are somehow diluted below a threshold level in the hybrids.     

Resistance of Trisomic and Diploid Hybrids of Beta vulgaris and B. procumbens to the Sugarbeet Nematode,Heterodera schachtiis Arnold

Trisomic and diploid hybrids of sugarbeet (Beta vulgaris L.) X wild beet (B. procumbens Chr. Sin.) inherited the gene for resistance to Heterodera schachtii Schm. from B. procumbens. The hybrids showed partial resistance to H. schachtii, manifested in failure of larvae to reach maturity. Although significantly greater numbers of female nematodes developed on plants inoculated with populations from the Netherlands or Italy than on plants inoculated with a population from the Salinas Valley, California. the totals for all populations on resistant plants were small. Greater numbers of males than females developed on root-slice cultures of resistant hyhrids when compared to a susceptible cultivar.     

Variations in the resistance to Heterodera rostochiensis of potatoes derived from Solarium tuberosum ssp. andigena and S. multidissectum

The proportion of larvae from ten populations of Heterodera rostochiensis Woll. that became female was determined on five potato clones containing genes for resistance derived from S. tuberosum ssp. andigena., on three with genes from S. multidissectum and on four with genes for resistance from both sources. Variations in the resistance of the clones bred from andigena, especially to two of the populations, suggest the presence of more than one gene for resistance.     

Mutual interference in the mass-reared fruit fly parasitoid,Diachasmimorpha longicaudata (Hymenoptera: Braconidae)

Diachasmimorpha longicaudata is a solitary endoparasitoid produced in Mexico for the biological control of Anastrepha fruit flies (Diptera: Tephritidae). We studied the effect of mutual interference among conspecific foraging females to better understand the parasitoid-host dynamics established in the mass-rearing system of this species. We used a constant host availability of 60 third instar larvae of Anastrepha ludens (per oviposition unit type Petri dish) that were individually exposed for 3 h to 1, 2, 4, 8, 10, 14, and 20 D. longicaudata females, seven days old without oviposition experience. The following parameters were evaluated: total number of attacked hosts (i.e., larvae with at least one oviposition scar), number of attacked hosts per female, adult emergence percentage, and female proportion. We also performed a second assay at mass-rearing level, contrasting the current proportion used in mass-rearing (~?two host larvae per female) with the alternative treatments (5 and 7.5 host larvae per female). Results showed that the density of females foraging on a patch exhibits an inverse relationship to the number of hosts attacked by one female. The highest values for adult emergence (73.4%), patch exploitation (94.6%), and female proportion (0.86) were obtained in the treatments with 7.5, 6, and 3 host larvae per female, respectively. Under mass-rearing conditions, we found that a lower density of females per cage (~?five larvae per female) reduced superparasitism levels without affecting adult emergence and the proportion of females. This suggests that the mass-rearing efficiency of D. longicaudata could be improved by reducing the density of foraging females.

     

Development, survival and fecundity of the urine fly, Scatella (Teichomyza) fusca and predation by the black dumpfly, Hydrotaea aenescens

Development, survival and fecundity for Scatella (Teichomyza) fusca Maquart (Diptera: Ephydridae) were studied at 20 ± 1 °C and 85 ± 10% r.h. Mean (± S.E.) developmental times of eggs, larvae and pupae were 4.0 ± 0.06, 14.9 ± 0.19 and 14.6 ± 0.11 days, respectively, the mean (± S.E.) survival of the original egg cohort to the start of larval, pupal and adult stage being 77.2 ± 3.2%, 54.5 ± 2.6% and 47.9 ± 3.5%, respectively. Females and males displayed approximately straight survivorship curves during adult life, implying constant mortality rates. Mean (± S.E.) adult longevity was 41.6 ± 2.98 days for females and 51.2 ± 3.91 days for males. Assuming a stable age distribution the population consisted of 56% eggs, 31% larvae, 6% pupae and 7% adults. Oviposition peaked when females were 25 days old, and the highest reproductive values (RV_x) (mean ± S.E.) ranged from 129.1 ± 7.57 to 138.5 ± 6.83 for individuals 17–27 days old. A female deposited a mean (± S.E.) of 614.7 ± 35.9 female eggs over a maximum life span of 93 days. The basic reproductive rate (R ₀) (mean ± S.E.) was 173.0 ± 14.2 female offspring per female and the intrinsic rate of natural increase of female individuals (r) (mean ± S.E.) was 0.088 ± 0.001 day^-1. The mean (± S.E.) generation time (T) was 57.8 ± 0.78 days. In cultures with equal numbers of first instar S. fusca larvae and predacious third instar larvae of Hydrotaea aenescens (synonymous Ophyra aenescens) Wiedemann, mean (± S.E.) survival to the adult stage of S. fusca (16.7 ± 8.8%) was significantly lower than in controls with S. fusca alone (58.3 ± 7.4%). The potential significance of predation by H. aenescens on S. fusca in pig farms is discussed.     

Butterfly oviposition preference is not related to larval performance on a polyploid herb

The preference–performance hypothesis predicts that female insects maximize their fitness by utilizing host plants which are associated with high larval performance. Still, studies with several insect species have failed to find a positive correlation between oviposition preference and larval performance. In the present study, we experimentally investigated the relationship between oviposition preferences and larval performance in the butterfly Anthocharis cardamines. Preferences were assessed using both cage experiments and field data on the proportion of host plant individuals utilized in natural populations. Larval performance was experimentally investigated using larvae descending from 419 oviposition events by 21 females on plants from 51 populations of two ploidy types of the perennial herb Cardamine pratensis. Neither ploidy type nor population identity influenced egg survival or larval development, but increased plant inflorescence size resulted in a larger final larval size. There was no correlation between female oviposition preference and egg survival or larval development under controlled conditions. Moreover, variation in larval performance among populations under controlled conditions was not correlated with the proportion of host plants utilized in the field. Lastly, first instar larvae added to plants rejected for oviposition by butterfly females during the preference experiment performed equally well as larvae growing on plants chosen for oviposition. The lack of a correlation between larval performance and oviposition preference for A. cardamines under both experimental and natural settings suggests that female host choice does not maximize the fitness of the individual offspring.     

Hybridisations between Mytilus edulis and Mytilus galloprovincialis and performance of pure species and hybrid veliger larvae at different temperatures

The mussels Mytilus edulis L. and Mytilus galloprovincialis Lamark hybridise naturally in the wild along the Atlantic coast of Europe producing a patchwork of mixed pure species and hybrid populations. Individuals of both species were spawned in the laboratory and were hybridised in a series of reciprocal crosses. After 72 h, the proportion of eggs which developed into larvae (%yield) and the proportion of those larvae which had a normal veliger morphology (%normality) were estimated and compared between pure species and hybrid families. There were no significant differences in %yield or %normality between pure species and hybrids, but significant differences were evident between the offspring from different parents irrespective of whether they were hybrids or pure species. Therefore confirmation of hybrid heterosis in laboratory studies should not be based on a single, or a few reciprocal crosses. Hybrid and pure species veliger larvae were grown for approximately 4 weeks at 10, 14 or 20 °C. In all trials, pure M. galloprovincialis larvae grew significantly faster at 20 °C than either reciprocal hybrid or pure M. edulis larvae. Irrespective of temperature, in general, hybrid larvae grew slower than larvae of either pure species. Increased exposure to planktonic predation due to slow growth can be interpreted as selection against hybrids and this may play a role in the structure and distribution of mixed pure species and hybrid populations.     

Interspecific hybrids between kenaf ( Hibiscus cannabinus ) and roselle ( H. sabdariffa )

The successful interspecific cross is reported for the first time between kenaf (Hibiscus cannabinus L.), a diploid species (2n=36) and roselle (Hibiscus sabdariffa L.), a tetraploid species (2n=72). Kenaf, grown for its bast fiber and also under investigation as a source of paper pulp, is fast-growing and well adapted to mechanical harvesting, but susceptible to root-knot nematodes. Roselle, also grown for its bast fiber, is slower growing, not well adapted to mechanical harvesting, but certain varieties are resistant to root-knot nematodes. Five hybrid plants were produced from the pollination of 4,445 flowers of kenaf with pollen from roselle; no hybrid plants were produced from 2,655 pollinations made in the reciprocal direction. One line of roselle was the parent of 3 of the 5 hybrids; one line of kenaf was the parent of 2 of these 3. The F₁ hybrids were triploid, and varied in vigor, growth habit and vegetative morphology, but had similar flowers. Two of the F₁ hybrids showed high pollen fertility, apparently as a result of restitution at first meiotic division leading to unreduced spores. These two hybrids each produced a small amount of seed, which gave rise to an F₂ population of 22 plants. The F₂ plants vary in vigor but are morphologically uniform, have thick leaves with mosaic sectors, and are presumably spontaneous allohexaploids. The theoretical possibilities of increasing the percentage of recovery of the F₁ interspecific hybrids and of developing a synthesized hybrid variety useful for bast fiber and paper pulp are discussed. Research done cooperatively by Crops Research Division and Agricultural Engineering Research Division, Agricultural Research Service, U.S. Department of Agriculture (USDA), and the University of Florida Agricultural Experiment Station. Cytological investigations were carried out at the Department of Biological Sciences, Florida State University. Thanks are extended to the many individuals and organizations who supplied material for this study, and to M. Griffin Bell and E. Otho Richey, Agricultural Research Technicians, for their enthusiastic assistance.     

Genetic variation for sexual dimorphism in flower size within and between populations of gynodioecious Thymus vulgaris

There has been very little empirical study of quantitative genetic variation in flower size in sexually dimorphic plant species, despite the frequent occurrence of flower size differences between sexual phenotypes. In this study we quantify the nature of quantitative flower size variation in females and hermaphrodites of gynodioecious Thymus vulgaris. In a field study, females had significantly smaller flowers than hermaphrodites, and the degree of flower size dimorphism varied significantly among populations. To quantify the genetic basis of flower size variation we sampled maternal progeny from 10 F₀ females in three populations (across the range of variation in flower size in the field), performed controlled crosses on F₁ offspring in the glasshouse and grew F₂ progeny to flowering in uniform field conditions. A significant population * sex interaction was again observed, hence the degree of sexual dimorphism shows genetic variation among populations. A significant family * sex interaction was also observed, indicating that the degree of sexual dimorphism shows genetic variation among families. Females showed significantly greater variation among populations and among families than hermaphrodites. Female flower size varied significantly depending on the degree of stamen abortion, with morphologically intermediate females having flowers more similar to hermaphrodites than to other females. The frequency of female types that differ in the degree of stamen abortion varied among populations and families and mean family female flower size increased as the proportion of intermediate female types increased across families. Variation in the degree of flower size dimorphism thus appears to be a result of variation in the degree of stamen abortion in females, the potential causes of which are discussed.