MORPHOLOGY,INHERITANCE, AND EVOLUTIONARY SIGNIFICANCE OF SEX REVERSAL IN TRIPSACUM DACTYLOIDES (POACEAE)

Inflorescences of Tripsacum dactyloides (L.) L. (Andropogoneae) are characterized by single female spikelets at one to several basal nodes and paired male spikelets at several nodes above them on each raceme. Female spikelets are one-flowered and male spikelets are two-flowered. A sex form variant was found in a wild population in north central Kansas and classified as T. dactyloides (L.) L. forma prolificum Dayton et Dewald. The variant of this native distant relative of maize (Zea mays L. spp. mays) differs from the normal form by having both pistillate and perfect rather than staminate spikelets in the terminal (tassel) portion of the inflorescence and by having two functional pistillate florets in the basal spikelets instead of one. A recessive major gene at a single locus regulates the change of the inflorescence from monoecious to gynomonoecious.     

LIFE-HISTORY STRATEGIES AND SEX RATIOS FOR A CULTIVAR AND A WILD POPULATION OF BUCHLOE DACTYLOIDES (GRAMINEAE)

Buchloe dactyloides (Nutt.) Engelm. is a perennial, sod-forming shortgrass of the grasslands of central North America from Montana to northern Mexico. The objectives of this research were to compare a cultivar (Sharps Improved) and a wild population from the Oklahoma panhandle, both in the field and in the greenhouse over several growing seasons, in regard to life-history strategies, sex ratios, and constancy of sex expression. Sex ratios of offspring from individual burs (usually 1–5 seeds per bur) and for all surviving plants germinated from seeds in the greenhouse were markedly different for the two populations. The wild population exhibited a 1: 1 male/female ratio with no monoecious plants and complete stability in sex expression over 3 yr of study in the greenhouse. In contrast, 13% of the cultivar plants were monoecious, and its unisexual plants showed a female bias. Burs of the two populations, in flats exposed to three separated month-long periods of favorable moisture with intervening “drought” periods, showed differential germinability of seeds within burs. The cultivar had less dormancy and produced a greater number of seedlings in the first and second periods; wild burs significantly exceeded cultivar burs in the number of seedlings during the third period and in the number of burs giving rise to seedlings in all three periods. Field and greenhouse measurements of vegetative (VRE) and sexual reproductive effort (SRE) per plant indicated that SRE was minimal (ca. 2% of aboveground biomass) in relation to asexual ramet propagation by rapidly growing stolons (VRE = ca. 80%). Mean SRE and VRE were not only similar for the populations, but also for the males and females within a population, although the values for individual plants were highly variable. It is concluded that Buchloe dactyloides is largely dioecious, with at least some wild populations showing a 1: 1 male/female ratio and constancy of sex expression, and that the bur as a dispersal unit containing 1–5 seeds is particularly appropriate for the breeding system and habitat ecology of Buchloe dactyloides.     

RELATIONSHIP BETWEEN SYNAPTOSPERMY AND DIOECY IN THE LIFE HISTORY STRATEGIES OF BUCHLOE DACTYLOIDES (GRAMINEAE)

Buchloe dactyloides (Nutt.) Engelm. is a perennial dioecious grass, exhibiting synaptospermy in that the 1–5 seeds in the dispersal unit (bur) are not released prior to germination. A major positive effect is that at least 50% of the burs contain both a male and female, making sexual reproduction possible and reducing founder effects upon colonization by one or a few burs. The objectives of this research were to examine the relationships between synaptospermy and dioecy in the life history strategies of Buchloe dactyloides, and to investigate whether the potential costs of synaptospermy would exceed its benefits if the species were not dioecious. An experiment that tested the effects of tightly clumping three seedlings vs. a uniform spacing of single seedlings at two densities and two moisture levels indicated only slightly negative effects of this aspect of synaptospermy. Although number of tillers and shoot biomass per plant were lower for clumped seedlings at all density and moisture combinations, only 50% of the possible uniform vs. clumped comparisons were significantly different. In addition, these effects, even at low moisture and/or high density, were not great enough to affect survivorship during the 16-week growth period. Experiments under both optimal and low moisture conditions showed that seeds within burs germinated at a significantly higher percentage than isolated or “hulled” seeds; in contrast to the differential timing of germination from burs, almost all of the seedlings from hulled seeds were produced during the first of three germination periods. Although survivorship for these seedlings was quite high under optimal conditions, it was greatly reduced under low moisture, being only one-fourth of that for seedlings arising from burs. The possible reduction in the wind dispersal of propagules due to their teeth and to their greater weight and size appears offset by more effective ungulate and water dispersal. Considering the evidence that the bur anchors the seedling in the soil, protects the seeds from fire, reduces precocious germination by requiring considerable moisture for saturation, and enhances seed longevity, the benefits of synaptospermy appear to outweigh the costs, even if those benefits related to dioecy are not considered.     

EVOLUTION OF DIOECY IN BUCHLOE DACTYLOIDES (GRAMINEAE): TESTS FOR SEX-SPECIFIC VEGETATIVE CHARACTERS,ECOLOGICAL DIFFERENCES,AND SEXUAL NICHE-PARTITIONING

Buchloe dactyloides is a perennial dioecious grass in which male and female inflorescences are so strikingly dimorphic that they were originally assigned to different genera. The objective of this paper is to present the results of tests for sex-specific vegetative characters, ecological differences, and sexual niche-partitioning, combining them with prior information on the reproductive biology of Buchloe for an evaluation of the key factors leading to the evolution of dioecy and sexual dimorphism. Field and greenhouse data were collected from Oklahoma and Kansas populations on vegetative characters, allocation to reproduction, and relative growth and competitive success along resource gradients. Except for greater susceptibility to leaf rust by males, there were no significant differences between males and females in vegetative characters, total biomass, or reproductive effort. Field studies of spatial distributions of males and females failed to show any relation to soil, topography, or soil moisture. In a 45-month greenhouse experiment starting at the seedling stage, the relative growth and competitive success of randomly paired individuals showed no evidence for differential competitive success or for niche-partitioning of males and females. The “outcrossing advantage” and subsequent sexual specialization of the female inflorescence appear to be the major factors underlying this dimorphic system.     

INHERITANCE OF SEX IN ACHLYA BISEXUALIS

Parental crosses, backcrosses, repeated backcrosses, and sib matings with several strains of Achlya bisexualis Coker and Couch have been made. Progeny were tested for sexual reaction with the parental strains. The distribution of sex observed could not be explained on the basis of any simple gene or chromosomal mechanism. Two factors were suggested which may affect the distribution of sex. They are selfing in the female and the complexity of the hormonal system which controls the expression of sex. The inheritance of a cycloheximide-resistant mutant argues against selfing; however, further studies with other genetic markers are needed.     

PROGENY SEX RATIOS IN DIOECIOUS SILENE LATIFOLIA (CARYOPHYLLACEAE)

Most sex ratios reported for Silene latifolia are female biased. As a result of experiments performed by Correns in the early 1900s, pollen tube competition has generally been accepted as the primary cause of these skewed ratios. We did four sets of hand pollinations in which we varied the size of pollen loads and placement of pollen along the filamentous stigma. The effect of pollen load size on progeny sex ratios was not statistically significant. Of 32 maternal families, 17 contained more females than males (one ratio deviated statistically from 1:1), and 13 contained more males than females. Paternal families exhibited a greater range of sex ratios, including three with a significant female bias and one with a significant male bias. Within experiments, neither the maternal parent nor where pollen was placed had a statistically significant effect on progeny sex ratios; the paternal effect was significant in one experiment. We suggest that sex ratios in Silene latifolia are not necessarily affected by the level of pollen competition. Other factors, including variation among males and sex-linked mortality, may help explain the skewed sex ratios that characterize populations of this species. Further, Correns' observations of excess females may have resulted from his use of interspecific hybrids.     

A MARKER-BASED METHOD FOR INFERENCES ABOUT QUANTITATIVE INHERITANCE IN NATURAL POPULATIONS

A marker-based method for studying quantitative genetic characters in natural populations is presented and evaluated. The method involves regressing quantitative trait similarity on marker-estimated relatedness between individuals. A procedure is first given for estimating the narrow sense heritability and additive genetic correlations among traits, incorporating shared environments. Estimation of the actual variance of relatedness is required for heritability, but not for genetic correlations. The approach is then extended to include isolation by distance of environments, dominance, and shared levels of inbreeding. Investigations of statistical properties show that good estimates do not require great marker polymorphism, but rather require significant variation of actual relatedness; optimal allocation generally favors sampling many individuals at the expense of assaying fewer marker loci; when relatedness declines with physical distance, it is optimal to restrict comparisons to within a certain distance; the power to estimate shared environments and inbreeding effects is reasonable, but estimates of dominance variance may be difficult under certain patterns of relationship; and any linkage of markers to quantitative trait loci does not cause significant problems. This marker-based method makes possible studies with long-lived organisms or with organisms difficult to culture, and opens the possibility that quantitative trait expression in natural environments can be analyzed in an unmanipulative way.     

SEX RATIO SELECTION AND THE EVOLUTION OF ENVIRONMENTAL SEX DETERMINATION IN LABORATORY POPULATIONS OF MENIDIA MENIDIA

What happens when a population with environmental sex determination (ESD) experiences a change to an extreme environment that causes a highly unbalanced sex ratio? Theory predicts that frequency-dependent selection would increase the proportion of the minority sex and decrease the level of ESD in subsequent generations. We empirically modeled this process by maintaining five laboratory populations of a fish with temperature-dependent sex determination (the Atlantic silverside, Menidia menidia) in extreme constant temperature environments that caused highly skewed sex ratios to occur initially. Increases in the minority sex consistently occurred from one generation to the next across all five populations, first establishing and then maintaining a balanced sex ratio until termination of the experiment at 8 to 10 generations. The extent to which the level of ESD changed as balanced sex ratios evolved, however, was not consistent. Two populations that experienced high temperatures each generation displayed a loss of ESD, and in one of these ESD was virtually eliminated. This suggests that temperature-insensitive, sex-determining genes were being selected. In populations maintained in low temperature environments, however, the level of ESD did not decline. Instead, the response of sex ratio to temperature was adjusted upward or downward, perhaps by selection of sex-determining genes sensitive to higher (or lower) temperatures. The two different outcomes at low versus high temperatures occurred independent of the geographic origin of the founding population. Our results demonstrate that ESD is capable of evolving in response to selection.     

DEVELOPMENT OF VOCALISATIONS IN NESTLING AND FLEDGLING HOUSE WRENS IN NATURAL POPULATIONS

ABSTRACT

We conducted a field study of the ontogeny of vocal signals in the house wren Troglodytes aedon during the nestling and fledgling phases of life. We did spectrographic analyses and quantification of the developmental changes that occurred in the acoustic features of the vocalisations. Evidence of progress to adult-like vocal patterns was of two types. First, nestling calls changed into a harsh-sounding call that resembles the adult chatter call, functionally a warning call. Second, fledglings also uttered subsong, and these vocalisations were similar to notes typical of adult male song. When the vocalisations produced by developing young were broken down into their constituent vocal features, we found that the time course of development was not strictly linear. Instead of a unidirectional change through the course of nestling and fledgling life, the trajectories of the vocal features fluctuated through time and sometimes exhibited abrupt changes. These sudden shifts occurred during nestling life as well as at the time of fledging. We speculate on the possible causes of these abrupt transitions. Changes in acoustic features upon fledging appear to be linked to new social functions.     

THE DISTRIBUTION OF SOLUBLE AND MEMBRANE-BOUND FORMS OF GLUTAMINASE IN PIG BRAIN

Abstract— About 10% of the glutaminase activity associated with pig brain mitochondria was readily extractable by a variety of techniques but the remainder was very resistant to extraction. These two forms, which have been termed the soluble and membrane-bound forms respectively, have been shown to differ in their responses to activation by phosphate and phosphate-borate containing buffers. Submitochondrial fractionation studies indicated that the soluble form was located in the mitochondrial inner matrix whereas the membrane-bound form was associated with the inner membrane. The mitochondria associated with the synaptosomes were found to contain only the membrane-bound form of the enzyme whereas both forms were present in the free brain mitochondria.     

MORPHOLOGICAL AND CYTOLOGICAL VARIABILITY IN TRIPSACUM DACTYLOIDES (GRAMINEAE)

A total of 235 accessions of Tripsacum dactyloides (L.) L. representing natural populations from 88 locations distributed throughout ten states of the United States were collected in the wild and studied morphologically and cytologically. Multivariate analysis of variance (MANOVA) was applied to 17 morphological characters recorded for 868 specimens from the 88 locations, which were further combined into 25 groups to facilitate analysis. Multivariate F ratios for six comparisons involving different groups were highly significant, indicating that the populations were significantly different on the basis of all 17 characters. Non-significant MANOVA F ratios for two comparisons between sympatric diploid and polyploid accessions showed them to be morphologically similar. Discriminant function (DF) analysis was employed to determine which characters contributed towards group discrimination. No one variable was found to distinguish between groups; rather, a combination of characters was necessary, and the combination changed with each comparison. Accessions were mainly either diploid (2n = 36) or tetraploid (2n = 72), with two triploids (2n = 54) reported from separate locations in Texas. Cytological data revealed that diploids and tetraploids occupy a more sympatric distribution than previously supposed. Morphological analysis suggested that diploids could not always be distinguished from polyploids on the basis of morphology alone, unless the appropriate characters were used. Information from cytological and morphological data showed Tripsacum dactyloides to be composed of many populations covering a wide range of variability, with each population possessing a characteristic combination of morphological attributes. It was concluded that T. dactyloides can be most meaningfully analyzed in terms of particular geographic regions, rather than attempting to elucidate natural relationships by correlating data from the entire United States.