Patterns of genetic diversity in outcrossing and selfing populations of Arabidopsis lyrata

Arabidopsis lyrata is normally considered an obligately outcrossing species with a strong self-incompatibility system, but a shift in mating system towards inbreeding has been found in some North American populations (subspecies A. lyrata ssp. lyrata). This study provides a survey of the Great Lakes region of Canada to determine the extent of this mating system variation and how outcrossing rates are related to current population density, geographical distribution, and genetic diversity. Based on variation at microsatellite markers (progeny arrays to estimate multilocus outcrossing rates and population samples to estimate diversity measures) and controlled greenhouse pollinations, populations can be divided into two groups: (i) group A, consisting of individuals capable of setting selfed seed (including autogamous fruit set in the absence of pollinators), showing depressed outcrossing rates (T(m) = 0.2-0.6), heterozygosity (H(O) = 0.02-0.06) and genetic diversity (H(E) = 0.08-0.10); and (ii) group B, consisting of individuals that are predominantly self-incompatible (T(m) > 0.8), require pollinators for seeds set, and showing higher levels of heterozygosity (H(O) = 0.13-0.31) and diversity (H(E) = 0.19-0.410). Current population density is not related to the shift in mating system but does vary with latitude. Restricted gene flow among populations was evident among all but two populations (F(ST) = 0.11-0.8). Group A populations were more differentiated from one another (F(ST) = 0.78) than they were from group B populations (F(ST) = 0.59), with 41% of the variation partitioned within populations, 47% between populations, and 12% between groups. No significant relationship was found between genetic and geographical distance. Results are discussed in the context of possible postglacial expansion scenarios in relation to loss of self-incompatibility.     

Do inbreeding depression and relative male fitness explain the maintenance of androdioecy in white mangrove, Laguncularia racemosa (Combretaceae)?

Mathematical models predict that to maintain androdioecious populations, males must have at least twice the fitness of male function in hermaphrodites. To understand how androdioecy is maintained in Laguncularia racemosa (white mangrove), outcrossing, inbreeding depression, and relative male fitness were estimated in two androdioecious populations and one hermaphroditic population. Outcrossing was estimated based on length of pollinator foraging bout and pollen carryover assumptions. Inbreeding depression was measured at three life stages: fruit set, seedling emergence, and seedling survivorship. The relative fitnesses of males and the male component of hermaphrodites were compared at these three stages and at the pollen production stage. Male frequency predictions generated by Lloyd's model were compared with observed frequencies in two androdioecious subpopulations. Outcrossing estimates were moderate for all populations (0.29-0.66). Inbreeding depression varied among populations (-0.03-0.86), but the strength of inbreeding depression did not increase with male frequency. Males produced significantly more flowers/inflorescence than hermaphrodites, but pollen production/flower did not differ. Male and hermaphroditic progeny did not differ significantly at other life stages. Populations of white mangrove with male plants were functionally androdioecious. Lloyd's model accurately predicted male frequency in one androdioecious subpopulation, but underestimated male frequency in the second subpopulation.     

Evolution of outcrossing in experimental populations of Caenorhabditis elegans

Caenorhabditis elegans can reproduce exclusively by self-fertilization. Yet, males can be maintained in laboratory populations, a phenomenon that continues to puzzle biologists. In this study we evaluated the role of males in facilitating adaptation to novel environments. For this, we contrasted the evolution of a fitness component exclusive to outcrossing in experimental populations of different mating systems. We introgressed a modifier of outcrossing into a hybrid population derived from several wild isolates to transform the wild-type androdioecious mating system into a dioecious mating system. By genotyping 375 single-nucleotide polymorphisms we show that the two populations had similar standing genetic diversity available for adaptation, despite the occurrence of selection during their derivation. We then performed replicated experimental evolution under the two mating systems from starting conditions of either high or low levels of diversity, under defined environmental conditions of discrete non-overlapping generations, constant density at high population sizes (N = 10⁴), no obvious spatial structure and abundant food resources. During 100 generations measurements of sex ratios and male competitive performance showed: 1) adaptation to the novel environment; 2) directional selection on male frequency under androdioecy; 3) optimal outcrossing rates of 0.5 under androdioecy; 4) the existence of initial inbreeding depression; and finally 5) that the strength of directional selection on male competitive performance does not depend on male frequencies. Taken together, these results suggest that androdioecious males are maintained at intermediate frequencies because outcrossing is adaptive.     

Pollen Production in the Androdioecious Datisca glomerata (Datiscaceae): Implications for Breeding System Equilibrium

Abstract Androdioecy (the presence of males and hermaphrodites in a breeding population) is a rare reproductive system in plants, with Datisca glomerata (Datiscaceae) representing the only well-documented example. Recent reports of high outcrossing rates, inbreeding depression, and high male pollen production satisfy theoretical predictions for the continued maintenance of androdioecy in populations of this species. However, in prior studies pollen production was measured indirectly in terms of numbers of anthers per flower—based on the assumption that male and hermaphroditic plant have equal numbers of flowers and that anthers from the two sexual morphs produce equivalent amounts of pollen. Herein, we demonstrate that male and hermaphrodite plants do not differ significantly in terms of flower number, but that pollen production in anthers from hermaphroditic plants is 12.6% higher than in anthers from male plants, thus refining the estimate of relative pollen fecundity of male versus hermaphrodite plants. The differential lowers the frequency of males predicted by theory, but is still consistent with the maintenance of androdioecy in this species.     

Androdioecy in the entomophilous tree Fraxinus ornus (Oleaceae)

Androdioecy (the coexistence of two genders, cosexuals and males, in a single population) is a rare breeding system. In terms of functional gamete production, androdioecy has been reported in a small number of wind-pollinated and insect-pollinated species. In this study we examine the floral biology, stability of gender, and fruit and seed production after self-pollination and outcrossing pollination in a potentially androdioecious tree, Fraxinus ornus , in southern France. Individual trees have either hermaphrodite flowers or male flowers, the latter lacking a well-developed gynoecium. The two genders produce morphologically similar pollen and were present in 1:1 or slightly male-biased ratios. Only hermaphrodites set fruit during 2–4 yr of observation at four different sites in southern France. Experimental pollinations of eight trees in two different sites showed that hermaphrodites produce viable pollen in dehiscent anthers and viable seeds. Thus, in terms of functional gamete production, F. ornus is a new case of functional androdioecy. Seven hermaphrodites were self-compatible, and such self-compatibility of hermaphrodites might improve the colonizing ability of F. ornus in a region where this species is actively expanding its range.     

Exploring directionality in spontaneous heart period and systolic pressure variability interactions in humans: implications in the evaluation of baroreflex gain

Although in physiological conditions RR interval and systolic arterial pressure (SAP) are likely to interact in a closed loop, the traditional cross-spectral analysis cannot distinguish feedback (FB) from feedforward (FF) influences. In this study, a causal approach was applied for calculating the coherence from SAP to RR (K(s-r)) and from RR to SAP (K(r-s)) and the gain and phase of the baroreflex transfer function. The method was applied, compared with the noncausal one, to RR and SAP series taken from 15 healthy young subjects in the supine position and after passive head-up tilt. For the low frequency (0.04-0.15 Hz) spectral component, the enhanced FF coupling (K(r-s) = 0.59 +/- 0.21, significant in 14 subjects) and the blunted FB coupling (K(s-r) = 0.17 +/- 0.17, significant in 4 subjects) found at rest indicated the prevalence of nonbaroreflex mechanisms. The tilt maneuver recovered FB influences (K(s-r) = 0.47 +/- 0.16, significant in 14 subjects), which were stronger than FF interactions (K(s-r) = 0.34 +/- 0.19, significant in 9 subjects). At the respiratory frequency, the RR-SAP regulation was balanced at rest (K(s-r) = 0.30 +/- 0.18 and K(r-s) = 0.29 +/- 0.20, significant in 11 and 8 subjects) and shifted toward FB mechanisms after tilt (K(s-r) = 0.35 +/- 0.19 and K(r-s) = 0.19 +/- 0.11, significant in 14 and 8 subjects). The causal baroreflex gain estimates were always lower than the corresponding noncausal values and decreased significantly from rest to tilt in both frequency bands. The tilt-induced increase of the phase lag from SAP to RR suggested a shift from vagal to sympathetic modulation. Thus the importance of nonbaroreflex interactions pointed out the necessity of accounting for causality in the cross-spectral analysis of the interactions between cardiovascular variables in healthy humans.     

Widespread functional android in Mercurialis annua L. (Euphorbiaceae)

The widespread coexistence of male and monoecious (cosexual) plants in Spanish, Portuguese and Moroccan populations of Mercurialis annua , an annual wind-pollinated ruderal, represents an important case of functional androdioecy, a rare breeding system in plants and animals. In M. annua , both males and cosexes disperse fully competent pollen. Quantitative gender varies discontinuously between males and cosexes, with males producing a mean of 6.09 times as much pollen as cosexes. It appears that gender is determined by a simple developmental switch, with male and cosexual inflorescences differing markedly in morphology: staminate flowers are borne on erect peduncles in males and in tight spiral clusters around a subsessile pistillate flower in cosexes. Males do not differ from cosexes in their biomass, but they are significantly taller, principally as a result of their greater internode lengths. The cosexual inflorescence is strongly protogynous so that outcrossing is favoured in dense stands, but seed-set is assured in cosexes isolated from prospective mates because of their ability to self-fertilize. Males typically occur at frequencies of less than about 30% in androdioecious populations, in accordance with theoretical predictions for functional androdioecy. In the genus Mercurialis , dioecy is the ancestral condition and monoecy and androdioecy, which occur in polyploid populations of M. annua , are derived. I argue here that androdioecy is most likely to evolve in plants (1) from dioecy, (2) in wind-pollinated species, and (3) in species with a colonizing habit. These predictions are also consistent with the limited published data available for other species.     

脱落酸对低温下雷公藤幼苗光合作用及叶绿素荧光的影响

以1年生雷公藤扦插苗为试材,研究低温胁迫下不同浓度外源脱落酸（ABA,0、5、10、15、20、25 mg·L^-1)叶面喷施处理对雷公藤叶片光合作用及叶绿素荧光参数的影响.结果表明：喷施20 mg·L^-1的ABA能显著提高雷公藤幼苗的抗冷性,减缓低温下雷公藤叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(g_s)、胞间CO₂浓度(C_i)的下降幅度,提高幼苗叶片的光合能力.低温处理6 d后,随着ABA浓度上升,雷公藤叶片的初始荧光(F_o)下降,最大荧光(F_m)和PSII最大光化学效率(F_v/F_m)上升,PSII实际光化学量子产量(Φ_PSⅡ)、光化学猝灭系数(q_P)先下降后上升,而非光化学猝灭系数(q_N)呈下降－上升－下降趋势.P_n、g_s、q_P、F_m和F_v/F_m均在20 mg·L^-1ABA处理时达到峰值.不同浓度ABA的相对电子传递速率(rETR)随着光化光强度增加呈先上升后下降的趋势,当光化光强度(PAR)达到395 μmol·m^-2s^-1时,各处理的rETR达到最高值,其中25 mg·L^-1和20 mg·L^-1ABA处理分别比对照高17.1%和5.2%.雷公藤叶片Φ_PSⅡ的光响应曲线均随光化光强度升高而下降,q_N的光响应曲线则呈相反趋势.     

Voltage-dependent blockade of connexin40 gap junctions by spermine

The effects of spermine and spermidine, endogenous polyamines that block many forms of ion channels, were investigated in homotypic connexin (Cx)-40 gap junctions expressed in N2A cells. Spermine blocked up to 95% of I(j) through homotypic Cx40 gap junctions in a concentration- and transjunctional voltage (V(j))-dependent manner. V(j) was varied from 5 to 50 mV in 5-mV steps and the dissociation constants (K(m)) were determined from spermine concentrations ranging from 10 micro M to 2 mM. The K(m) values ranged from 4.9 mM to 107 micro M for 8.6 < or = V(j) < or = 37.7 mV, within the physiological range of intracellular spermine for V(j) > or = 20 mV. The K(m) values for spermidine were > or = 5 mM. Estimates of the electrical distance (delta) for spermine (z = +4) and spermidine (z = +3) were 0.96 and 0.76 respectively. Cx40 single channel conductance was 129 pS in the presence of 2-mM spermine and channel open probability was significantly reduced in a V(j)-dependent manner. Similar concentrations of spermine did not block I(j) through homotypic Cx43 gap junctions, indicating that spermine selectively blocks Cx40 gap junctions. This is contrary to our previous findings that large tetraalkylammonium ions, also known to block several forms of ion channels, block junctional currents (I(j)) through homotypic connexin Cx40 and Cx43 gap junctions.     

Genic diversity, genetic structure, and mating system of Brewer spruce (Pinaceae), a relict of the Arcto-Tertiary forest

Brewer spruce (Picea breweriana), a relict of the widespread Arcto-Tertiary forests, is now restricted to a highly fragmented range in the Klamath Region of California and Oregon. Expected heterozygosity for 26 isozyme loci, averaged over 10 populations, was 0.121. More notable than the relatively high level of diversity when compared to other woody endemics was the strong decrease in expected heterozygosity with latitude. Differentiation (F(ST)) was 0.152, higher than values for many north temperate conifers with larger distributions. The number of migrants per generation (Nm) was 1.34 or 2.70, depending on the method of estimation. Inbreeding appeared low; F(IS) was only 0.003, in agreement with multilocus population outcrossing rates (t(m)), which were generally well above 0.90. No difference in t(m) was found between isolated vs. clustered trees. However, the number of seeds per cone was greatest in the densest populations; t(m) is a measure of effective outcrossing after mortality in the embryonic stage, whereas a reduced number of seeds per cone indicates self pollination. Selfing increased after logging; outcrossing rate before logging was 0.961 and after logging, 0.756. Despite Brewer spruce's narrow, fragmented distribution, the outlook for its conservation was good, with the exception of possible negative effects of logging.     

Population genetic structure of the gynodioecious Thymus vulgaris L. (Labiatae) in southern France

In a self-compatible gynodioecious species, the abundance of female plants (which are obligate outcrossers) relative to hermaphrodites (which may self and outcross) may be a critical factor influencing genetic diversity and population structure. In the gynodioecious Thymus vulgaris L., female frequency varies from 5 to 95%, providing a suitable model to examine this issue. In this study, we use allozyme markers to (1) evaluate the relationship between female frequency, genetic diversity and population structure, (2) determine whether females and hermaphrodites vary in heterozygote deficiency and (3) examine whether other factors such as plant density are related to heterozygote deficiency. Twenty three natural populations, with female frequencies ranging from 11 to 92%, were sampled in and around the St-Martin-de-Londres basin in southern France. Based on four polymorphic allozyme loci, we found no significant correlation between female frequency and heterozygote deficiency. A significant (P < 0.05) F_IS value over loci and over populations of 0.11 was detected. The F_IS value per population showed a significant heterozygote deficiency in 11 of the 23 populations. However, no significant difference between female and hermaphrodite F_IS values was found. A significant heterozygote deficiency only occurred in populations of intermediate density. There was little differentiation among populations (F_ST = 0.038) nor among subpopulations within each population. The significant F_IS values are thus mostly due to inbreeding effects. The lack of a correlation between F_IS values and female frequency may be due to outcrossing in hermaphrodites and/or restoration of male fertility which may occur to a greater extent at low female frequency. The similarity of female and hermaphrodite F_IS values indicates that females may occasion high levels of biparental inbreeding.     

The genetic basis and experimental evolution of inbreeding depression in Caenorhabditis elegans

Determining the genetic basis of inbreeding depression is important for understanding the role of selection in the evolution of mixed breeding systems. Here, we investigate how androdioecy (a breeding system characterized by partial selfing and outcrossing) and dioecy (characterized by obligatory outcrossing) influence the experimental evolution of inbreeding depression in Caenorhabditis elegans. We derived inbred lines from ancestral and evolved populations and found that the dioecious lineages underwent more extinction than androdioecious lineages. For both breeding systems, however, there was selection during inbreeding because the diversity patterns of 337 single-nucleotide polymorphisms (SNPs) among surviving inbred lines deviated from neutral expectations. In parallel, we also followed the evolution of embryo to adult viability, which revealed similar starting levels of inbreeding depression in both breeding systems, but also outbreeding depression. Under androdioecy, diversity at a neutral subset of 134 SNPs correlated well with the viability trajectories, showing that the population genetic structure imposed by partial selfing affected the opportunity for different forms of selection. Our findings suggest that the interplay between the disruptions of coevolved sets of loci by outcrossing, the efficient purging of deleterious recessive alleles with selfing and overdominant selection with outcrossing can help explain mixed breeding systems.     

Inferring recent outcrossing rates using multilocus individual heterozygosity: application to evolving wheat populations

Using multilocus individual heterozygosity, a method is developed to estimate the outcrossing rates of a population over a few previous generations. Considering that individuals originate either from outcrossing or from n successive selfing generations from an outbred ancestor, a maximum-likelihood (ML) estimator is described that gives estimates of past outcrossing rates in terms of proportions of individuals with different n values. Heterozygosities at several unlinked codominant loci are used to assign n values to each individual. This method also allows a test of whether populations are in inbreeding equilibrium. The estimator's reliability was checked using simulations for different mating histories. We show that this ML estimator can provide estimates of outcrossing rates for the final generation outcrossing rate (t(0)) and a mean of the preceding rates (t(p)) and can detect major temporal variation in the mating system. The method is most efficient for low to intermediate outcrossing levels. Applied to nine populations of wheat, this method gave estimates of t(0) and t(p). These estimates confirmed the absence of outcrossing t(0) = 0 in the two populations subjected to manual selfing. For free-mating wheat populations, it detected lower final generation outcrossing rates t(0) = 0-0.06 than those expected from global heterozygosity t = 0.02-0.09. This estimator appears to be a new and efficient way to describe the multilocus heterozygosity of a population, complementary to Fis and progeny analysis approaches.     

不同浓度Ni、Cu处理对骆驼蓬光合作用和叶绿素荧光特性的影响水

以骆驼蓬幼苗为材料,采用盆栽试验研究不同浓度(0、50、100、200、400 mg·kg-1)Ni、Cu处理对骆驼蓬叶片光合作用、叶绿素荧光特性及生长状况的影响.结果表明:随着Ni浓度的增加,骆驼蓬幼苗叶片的光合色素含量、净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、PS Ⅱ最大光化学效率(Fv/Fm)、PS Ⅱ电子传递量子产率(φpsⅡ)、光化学猝灭系数(qp)及各项生长指标均呈显著下降趋势,而细胞间隙CO2浓度(Ci)和非光化学猝灭系数(qn)呈显著增加趋势,其中Pn的下降主要是由非气孔限制所致;骆驼蓬幼苗叶片的光合色素含量、Pn、Gs、Tr、Ci、Fv/Fm、φpsⅡ、qp及各项生长指标均在50 mg·kg-1Cu处理时达到峰值,叶绿素a和b、Pn、Gs、Tr、Ci、Fv/Fm及各项生长指标值在100 mg·kg-1Cu处理时仍微高于对照,而后随Cu浓度的增加,光合色素含量、Pn、Gs、Tr、Ci、Fv/Fm、φpsⅡ、qp及各项生长指标均呈下降趋势,qN呈增加趋势,其中Pn的下降主要是由气孔限制所致.     

Genetic diversity and the mating system of a rare Mexican pinon, Pinus pinceana, and a comparison with Pinus maximartinezii (Pinaceae)

Weeping pi?on (Pinus pinceana) has a restricted and fragmented range, trees are widely scattered within populations, and reproduction is limited. Nevertheless, genetic diversity was high; based on 27 isozyme loci in 18 enzyme systems, unbiased expected heterozygosity averaged 0.174. Differentiation also was high (F(ST) = 0.152), reflecting isolation between southern, central, and northern fragments of the range. Among populations in the northern fragment, F(ST) was only 0.056, and the number of migrants per generation (Nm) was 4.21, which should preclude fixation. Nm between central and southern populations or between them and populations in the northern fragment was lower, 0.99-1.66, indicating a degree of genetic isolation. Multilocus outcrossing rates (t(m)) ranged from 0.836 in the south to 0.897 in the north. Therefore, selfing is low but statistically significant. The equilibrium inbreeding coefficient (F(e)) calculated from t(m) was in good agreement with observed inbreeding coefficients, suggesting that weeping pi?on may be near equilibrium with respect to inbreeding and selection against selfed trees. Weeping pi?on was variable at all loci polymorphic in maxipi?on (Pinus maximartinezii) and, therefore, qualifies as a possible progenitor of maxipi?on. Because of the high level of diversity, reasonable levels of gene flow within the northern fragment of weeping pi?on's range, high rates of outcrossing, and, perhaps, only weak selection against inbred trees, protection in reserves would be a viable option for conservation.     

Gammaherpesviruses encode functional dihydrofolate reductase activity

We overexpressed and purified from Escherichia coli the dihydrofolate reductase (DHFR) of the gammaherpesviruses human herpesvirus 8 (HHV-8), herpesvirus saimiri (HVS), and rhesus rhadinovirus (RRV). All three enzymes proved catalytically active. The K(m) value of HHV-8 DHFR for dihydrofolate (DHF) was 2.02+/-0.44 microM, that of HVS DHFR was 4.31+/-0.56 microM, and that of RRV DHFR is 7.09+/-0.11 microM. These values are approximately 5-15-fold higher than the K(m) value reported for the human DHFR. The K(m) value of HHV-8 DHFR for NADPH was 1.31+/-0.23 microM, that of HVS DHFR was 3.78+/-0.61 microM, and that of RRV DHFR was 7.47+/-0.59 microM. These values are similar or slightly higher than the corresponding K(m) value of the human enzyme. Methotrexate, aminopterin, trimethoprim, pyrimethamine, and N(alpha)-(4-amino-4-deoxypteroyl)-N(delta)-hemiphthaloyl-L-ornithine (PT523), all well-known folate antagonists, inhibited the DHFR activity of the three gammaherpesviruses competitively with respect to DHF but proved markedly less inhibitory to the viral than towards the human enzyme.     

Human cathepsin F: expression in baculovirus system, characterization and inhibition by protein inhibitors

Recombinant full-length human procathepsin F, produced in the baculovirus expression system, was partially processed during the purification procedure to a form lacking the N-terminal cystatin-like domain and activated with pepsin. Active cathepsin F efficiently hydrolyzed Z-FR-MCA (kcat/Km=106 mM(-1) s(-1)) and Bz-FVR-MCA (kcat/Km=8 mM(-1) s(-1)), whereas hydrolysis of Z-RR-MCA was very slow (kcat/Km<0.2 mM(-1) s(-1)). Cathepsin F was rapidly and tightly inhibited by cystatin C, chicken cystatin and equistatin with Ki values in the subnanomolar range (0.03-0.47 nM), whereas L-kininogen was a less strong inhibitor of the enzyme (Ki=4.7 nM). Stefin A inhibited cathepsin F slowly (kass=1.6 x 10(5) M(-1) s(-1)) and with a lower affinity (Ki=25 nM). These data suggest that cathepsin F differs from other related endopeptidases by considerably weaker inhibition by stefins.     

Cost of flight in the zebra finch ( Taenopygia guttata): a novel approach based on elimination of (13)C labelled bicarbonate

On three separate occasions, five zebra finches ( Taenopygia guttata) were injected intraperitoneally with 0.2 ml 0.29 M NaH(13)CO(3)solution and placed immediately into respirometry chambers to explore the link between (13)C elimination and both O(2) consumption (VO(2)) and CO(2) production (VCO(2)). Isotope elimination was best modelled by a mono-exponential decay. The elimination rate (k(c)) of the (13)C isotope in breath was compared to VO(2) (ml O(2)/min) and VCO(2) (ml CO(2)/min) over sequential 5-min time intervals following administration of the isotope. Elimination rates measured 15-20 min after injection gave the closest relationships to VO(2) ( r(2) =0.82) and VCO(2) ( r(2)=0.63). Adding the bicarbonate pool size (N(c)) into the prediction did not improve the fit. A second group of birds ( n=11) were flown for 2 min (three times in ten birds and twice in one) between 15 min and 20 min following an injection of 0.2 ml of the same NaH(13)CO(3) solution. Breath samples, collected before and after flight, were used to calculate k(c) over the flight period, which was converted to VO(2) and VCO(2) using the equation generated in the validation experiment for the corresponding time period. The energy expenditure (watts) during flight was calculated from these values using the average RQ measured during flight of 0.79. The average flight cost measured using the bicarbonate technique was 2.24+/-0.11 W (mean+/-SE). This average flight cost did not differ significantly from predictions generated by an allometric equation formulated by Masman and Klaassen (1987 Auk 104:603-616). It was however substantially higher than the predictions based on the aerodynamic model of Pennycuick (1989 Oxford University Press), which assumes an efficiency of 0.23 for flight. The flight efficiency in these birds was 0.11 using this model. Flight cost was not related to within-individual variation [general linear model (GLM) F(1,31)=1.16, P=0.29] or across-individual variations in body mass (GLM F(1,31)=0.26, P=0.61), wingspan (regression F(1,10)=0.01, P=0.94) or wing loading (regression F(1, 31)=0.001, P=0.99) in this sample of birds.     

Plant regeneration from hypocotyl segments of Lupinus luteus cv. L. Aurea

Complete plants of Lupinus luteus L. cv. Aurea that were regenerated from hypocotyl segments, bloomed, produced seeds and were efficiently nodulated by Bradyrhizobium sp. strains. The highest rates of shoot formation were obtained on A medium plus 1.3% agar with 10.0 M 2-isopentenyladenine (2iP) and 0.11 M naphthaleneacetic acid (NAA); the best rooting was achieved on a medium with 0.5 M NAA plus 0.05 M 2iP. Afterward, plantlets were transferred to either perlite or peat-containing pots and irrigated with a N-free nutrient solution until maturity. Direct rooting of hypocotyls could also be obtained on A medium with 1% agar.     

Genetic variation in outcrossing rate and correlated floral traits in a population of grain amaranth (Amaranthus cruentus L.)

The genetic control of breeding system in a population of grain amaranth is described here by the estimates of outcrossing rate variation among individuals and their response to mass selection. In the first generation (S₀), a single morphological marker locus(R/r) controlling plant pigmentation was used for the estimation of outcrossing rate. A mixture of genotypes RR (red) and rr (green) mixed in equal proportions was grown and all rr plants were progeny-tested for estimating the outcrossing rate (t), using the relationship t=H/p, where H = proportion of heterozygotes (Rr) in the progeny, and p = frequency of alleleR. A full range (0 to 100%) of outcrossing rates was found in the first generation, % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGabeiDayaara% Gaeyypa0JaaGimaiaac6cacaaIZaGaaGymaiaacYcacqGHGaaicqGH% GaaicuaHdpWCgaqcamaaBaaaleaacaqG0baabeaakiabg2da9iaaic% dacaGGUaGaaGOmaiaaiwdaaaa!441B!\[{\text{\bar t}} = 0.31, \hat \sigma _{\text{t}} = 0.25\], and the distribution was significantly skewed toward high outcrossing.A bidirectional mass selection experiment was initiated for high and low outcrossing rates. In the Hi and Lo lines (Generation S₁), the rate of outcrossing was estimated using the R locus in addition to two other completely dominant morphological markers(B, G1). Relative sex ratios in the monoecious inflorescences of selected lines were estimated by the number of male flowers per glomerule. Outcrossing rate was significantly different between the Hi and Lo lines, and Lo line exhibited much higher male fertility than the Hi line. Such a response to selection indicated a polygenic control of sex expression, and consequently, of the potential for outcrossing in this population. In the second generation (S₂), selection was based on the number of male flowers/glomerule as an index of outcrossing ability. Differences in sex expression between the Hi and Lo lines in S₂ generation had 1.0 male flower/glomerule, while the Hi selection individuals had more or less rectangular distribution from 0.1 to 1.0 male flowers/glomerule.Thus, breeding system in amaranth, as represented by the sex ratios within glomerules and the outcrossing rate parameters, responded to bidirectional selection. Such a genetic component of variation in breeding system has significant implications in the evolution of amaranth species and landraces under domestication.