Evolutionary loss of sepals and/or petals in detarioid legume taxa Aphanocalyx, Brachystegia, and Monopetalanthus (Leguminosae: Caesalpinioideae)

Floral development using scanning electron microscopy is compared in several taxa of the Brachystegia subtribal group of caesalpinioid tribe Detariae. This group is characterized by missing sepals and/or petals. In Aphanocalyx djumaensis, Monopetalanthus durandii, and two Brachystegia species, one sepal is initiated in median abaxial position. In the first two, one or two additional sepal rudiments may initiate late. Brachystegia species have all five sepals, which remain scalelike. In Aphanocalyx and Monopetalanthus, one petal initiates adaxially and medianly (a position atypical for the first initiated petal in the family); additional petal rudiments may form in lateral sites. In Brachystegia, five petals are initiated unidirectionally on a meristem ring, but all are suppressed after initiation. In all taxa, ten stamens are initiated on a ring meristem: unidirectionally in Monopetalanthus, bidirectionally in Brachystegia, vs. in erratic order in Aphanocalyx. Carpel and petal initiation are concurrent. Different organ whorls overlap in time in Monopetalanthus and Brachystegia. In all, the floral apex characteristically is elongate radially and narrow tangentially after bracteole initiation. Two ontogenetic features, the meristem ring and the radially elongate post-bracteole floral apex, appear to be possible synapomorphies for the Brachystegia group.     

Andromonoecy in Solanum lycocarpum A. St. -Hil. (Solanaceae): Floral attributes,visitors and variation in sexual expression over time

Sexual expression in andromonoecious species—those in which a single individual can bear both staminate and hermaphroditic flowers—may vary among reproductive events in the same plant, among individuals and across populations. This variation influences, in turn, the individual contribution of hermaphroditic plants via male and female fitness functions (i.e., Lloydʼs phenotypic gender). However, temporal variation in sexual expression in andromonoecious species and its relationship with seasonal changes in climatic conditions remain poorly understood. Here we analyze floral attributes, visitors and variation in sexual expression in three populations of Solanum lycocarpum A. St. -Hil. Seasonality in the production of floral types, the mating system and floral visitors were also investigated. Hermaphroditic flowers produced more pollen grains, but the pollen of staminate flowers had higher viability. Only hermaphroditic flowers produced fruits, and ovules in staminate flowers were sterile. Solanum lycocarpum is mainly pollinated by large bees with the ability to vibrate flowers. Phenotypic gender varied throughout the year, and the seasonal production of staminate flowers is associated with the local climate. We suggest that the higher and seasonally variable relative abundance of staminate flowers compared to the low and uniform production of hermaphroditic flowers may be explained by (a) the very high energetic costs incurred in producing large fruits, which in turn make hermaphroditic flower production very costly, and (b) the potentially lower energy expenditure of the smaller staminate flowers with reduced pistils and non-viable ovules that allow them to rapidly respond to climate variability.     

Evolutionary rate variation in Old World monkeys

We analysed over 8 million base pairs of bacterial artificial chromosome-based sequence alignments of four Old World monkeys and the human genome. Our findings are as follows. (i) Genomic divergences among several Old World monkeys mirror those between well-studied hominoids. (ii) The X-chromosome evolves slower than autosomes, in accord with ‘male-driven evolution’. However, the degree of male mutation bias is lower in Old World monkeys than in hominoids. (iii) Evolutionary rates vary significantly between lineages. The baboon branch shows a particularly slow molecular evolution. Thus, lineage-specific evolutionary rate variation is a common theme of primate genome evolution. (iv) In contrast to the overall pattern, mutations originating from DNA methylation exhibit little variation between lineages. Our study illustrates the potential of primates as a model system to investigate genome evolution, in particular to elucidate molecular mechanisms of substitution rate variation.     

Development of clover yellow vein virus as an efficient, stable gene-expression system for legume species

A highly infectious cDNA clone of clover yellow vein virus (pClYVV) was tested as a viral vector, especially for legume species. The genes for green fluorescent protein (GFP) and soybean glutamine synthetase (GS) were inserted between the genes for P1 and HC-Pro on pClYVV to create three recombinant plasmids: pClYVV-GFP, pClYVV-GFP-GS, and pClYVV-GFP:GS. In the former two constructs all the junctions between the inserted proteins contained the sequences of protease cleavage recognition sites, whereas the third construct expressed a fusion of GFP and GS. Western blot analyses showed that GFP and GS appeared to have been precisely excised from the viral polyprotein with the viral proteases (P1 and NIa). Under UV irradiation, green fluorescence was detected in infected broad bean, kidney bean, and soybean plants. The stability of the constructs in the symptomatic tissues was confirmed by RT-PCR and Western blot analyses. The plants expressing GS together with GFP became tolerant to the herbicide glufosinate, and flowered early. As the GS gene, one of the nodulin genes for nitrogen fixation, is expressed in legume species, this system will be useful for examining the function of genes important to legume plants.     

Carboxylesterases from the seeds of an underutilized legume, Mucuna pruriens; isolation, purification and characterization

Two carboxylesterases (ME-III and ME-IV) have been purified to apparent homogeneity from the seeds of Mucuna pruriens employing ammonium sulfate fractionation, cation exchange chromatography on CM-cellulose, gel-permeation chromatography on Sephadex G-100 and preparative PAGE. The homogeneity of the purified preparations was confirmed by polyacrylamide gel electrophoresis (PAGE), gel-electrofocussing and SDS–PAGE. The molecular weights determined by gel-permeation chromatography on Sephadex G-200 were 20.89 kDa (ME-III) and 31.62 kDa (ME-IV). The molecular weights determined by SDS–PAGE both in the presence and absence of 2-mercaptoethanol were 21 kDa (ME-III) and 30.2 kDa (ME-IV) respectively, suggesting a monomeric structure for both the enzymes. The enzymes were found to have Stokes radius of 2.4 nm (ME-III) and 2.7 nm (ME-IV). The isoelectric pH values of the enzymes, ME-III and ME-IV, were 6.8 and 7.4, respectively. ME-III and ME-IV were classified as carboxylesterases employing PAGE in conjunction with substrate and inhibitor specificity. The K_m of ME-III and ME-IV with 1-naphthyl acetate as substrate was 0.1 and 0.166 mM while with 1-naphthyl propionate as substrate the K_m was 0.052 and 0.0454 mM, respectively. As the carbon chain length of the acyl group increased, the affinity of the substrate to the enzyme increased indicating hydrophobic nature of the acyl group binding site. The enzymes exhibited an optimum temperature of 45 °C (ME-III) and 37 °C (ME-IV), an optimum pH of 7.0 (ME-III) and 7.5 (ME-IV) and both the enzymes (ME-III and ME-IV) were stable up to 120 min at 35 °C. Both the enzymes were inhibited by organophosphates (dichlorvos and phosphamidon), but resistant towards carbamates (carbaryl and eserine sulfate) and sulphydryl inhibitors (p-chloromercuricbenzoate, PCMB).     

Characteristics of an ideotype acid tolerant pasture legume symbiosis in Mediterranean agriculture

A series of experiments has led to the following concept of eight characteristics being required in an acid-tolerant pasture legume symbiosis for use in ley-farming: In the bulk soil i -microsymbiont (preferably of the Bradyrhizobium genus) capable of maintaining high numbers into autumn, through processes which allow saprophytic function at low pH such as regulation of its internal pH, ii -microsymbiont with a carboxylated cell surface electrochemistry stable under the influence of ambient pH in its interactions with soil colloids, minerals and root surfaces. In the rhizosphere iii -microsymbiont capable of appreciable growth in response to substrate availability, iv icrosymbiont able to recognise root exudates allowing interaction with its nodD gene protein, v -microsymbiont whose nodABC gene products (nod metabolites) are pH stable and which induce cortical cell division and root-hair curling in the host, vi -microsymbiont whose surface polysaccharides and proteins are pH stable to allow attenuation of the cell at the root surface, At the root surface vii-host apoplast function unaffected by low pH such that it may (a) produce pH stable exudates capable of interacting with the rhizobial nodD gene protein and (b) receive rhizobial nod metabolites and respond physiologically, viii-root able to produce pH stable organic acids for linkage with rhizobial cell surface structures.These ideotype characteristics reflect our current understanding of the mechanisms of acid tolerance in the nodulation phase of species such as Medicago polymorpha and M. murex, and acid soil tolerance in rhizobial inoculant strains such as WSM540.     

Contrasting responses of legume versus non-legume shrubs to soil water and nutrient shortages in the Mu Us Sandland

Aims Legumes and non-legumes usually differ in using soil water and nutrients. Both water and nutrients are scarce in the semi-arid Mu Us Sandland where legume and/or non-legume shrubs coexist/dominate. Here, we addressed the responses of legume versus non-legume shrubs to different soil water and nutrient conditions.Methods We conducted an experiment in which a legume (Hedysarum laeve) and a non-legume (Artemisia ordosica) were used, both of which are dominant species in the Mu Us Sandland. Seedlings of these two species were subjected to three water levels (45.0, 67.5 and 90.0 ml every 3 days) and three nutrient treatments (0, 0.1% and 0.2% nutrient solution every week) during the experiment.Important findings Interactions between water and nutrients on total biomass, root weight ratio and rain use efficiency (RUE) were detected in A. ordosica but not in H. laeve, suggesting that water effects on A. ordosica but not on H. laeve are dependent on soil nutrients. Nutrient addition alleviated drought stress and increased RUE in A. ordosica. The interspecific differences in response to soil water and nutrients may be linked to the ability of plants to fix nitrogen. In addition, under low-soil water or nutrient conditions, H. laeve produced more biomass than A. ordosica, and the opposite was the case under high-soil resources. The relationship between relative growth rate (RGR) and RUE [or nutrient use efficiency (NUE)] varied with two species. RGR of A. ordosica was positively correlated with both RUE and NUE while RGR of H. laeve was negatively correlated with NUE. The different responses may be linked to the trade-off between high-growth rate and low-resource use efficiency.     

Increasing phosphorus concentration in seed of annual pasture legume species increases herbage and seed yields

In glasshouse experiments with low levels of soil applied phosphorus (P), yields of four annual pasture legumes (Medicago polymorpha, Trifolium subterraneum, T. balansae, Ornithopus compressus) increased with increasing P concentration in the seed. In a further experiment, M. polymorpha cv. Serena was grown at the same plant density from seed of two P concentrations and two seed sizes when two levels of finely ground superphosphate were applied to the soil. Higher P concentrations in the seed increased yields of dried tops by about 30% for the first harvest (21 days), 20% for the second harvest (52 days), and 9% at maturity (103 days), and seed yields by 11%. Larger seeds increased yields of dried tops by between 6–46% for the first two harvests but at maturity yields of dried tops and seed were unaffected by seed size. None of the interactions were statistically significant (P>0.05), except for the first harvest when two interactions (P concentration in the seed × seed size (i.e. P content in seed), and P applied to the soil × P concentration in the seed × seed size) were significant at P<0.05 level. In a field experiment, Trifolium subterraneum clover seed (two cultivars) of the same size but with two different P concentrations was sown at the same plant density and two levels of granulated (0.2–5 mm) superphosphate were applied to the soil surface. The higher level of superphosphate increased dried herbage yields of the dense clover swards by three- to four-fold 90 and 120 days after sowing. The higher P concentration in the seed increased yields of dried herbage by between 50 to 25%, depending on the level of P applied to the soil and the harvest date.     

Pollen and Resource Limitation in Veratrum nigrum L. （Liliaceae）, an Andromonoecious Herb

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman＇s principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions.     

Sex‐specific selection on plant architecture through “budget” and “direct” effects in experimental populations of the wind‐pollinated herb,Mercurialis annua

Sexual selection may contribute to the evolution of plant sexual dimorphism by favoring architectural traits in males that improve pollen dispersal to mates. In both sexes, larger individuals may be favored by allowing the allocation of more resources to gamete production (a “budget” effect of size). In wind‐pollinated plants, large size may also benefit males by allowing the liberation of pollen from a greater height, fostering its dispersal (a “direct” effect of size). To assess these effects and their implications for trait selection, we measured selection on plant morphology in both males and females of the wind‐pollinated dioecious herb Mercurialis annua in two separate experimental common gardens at contrasting density. In both gardens, selection strongly favored males that disperse their pollen further. Selection for pollen production was observed in the high‐density garden only, and was weak. In addition, male morphologies associated with increased mean pollen dispersal differed between the two gardens, as elongated branches were favored in the high‐density garden, whereas shorter plants with longer inflorescence stalks were favored in the low‐density garden. Larger females were selected in both gardens. Our results point to the importance of both a direct effect of selection on male traits that affect pollen dispersal, and, to a lesser extent, a budget effect of selection on pollen production.     

Lesions to the Medial Preoptic Area Affect Singing in the Male European Starling (Sturnus vulgaris)

The aromatization of testosterone (T) in the medial preoptic nucleus (POM) is known to regulate male courtship and sexual behaviors expressed prior to, and in anticipation of, copulation. Singing in male European starlings is used to attract mates prior to physical sexual contact, suggesting that the POM might be involved. The present study was performed to examine the effects of lesions targeting the POM on singing and courtship behavior in reproductively active male starlings. A significant decrease in song output and the gathering of green nest materials was observed in males with lesions to the POM compared to males with damage to brain areas outside of the POM. Lesions did not affect a male's tendency to remain near a female or to occupy a nestbox, suggesting that the effects of POM lesions were specific to courtship behaviors. Behavioral differences were not related to testis mass or volume, and GnRH immunoreactivity was observed within the hypothalamus and median eminence for each male, suggesting that the effects of POM lesions were related specifically to POM involvement in song expression rather than to a disruption of the GnRH axis. These results suggest a general role for the POM in the expression of behaviors related to sexual arousal or anticipation, including song.     

Mating system of an exceptional primate, the olive colobus (Procolobus verus)

In the olive colobus (Procolobus verus), many groups have multiple males and the males have large testes. This indicates that even though this species lives in small groups, single males do not monopolize the groups. We investigated the strategies employed by males to secure their mating success, and sought to determine whether the lack of male monopolization was a result of female mating strategies, as indicated by the exaggerated sexual swellings of the females. Four study groups were monitored for demographic changes, and group composition was determined in six additional groups in Taï National Park, Ivory Coast, between 1994 and 1999. Social behavior was recorded by scan and focal sampling in the study groups. The almost permanent association of olive colobus with Diana monkeys (Cercopithecus diana) in effect provided males a resource at which they could expect females to visit and sometimes even permanently join them, as well as protection from predators. As alternative strategies for obtaining females, one male took over the group of another male and one male immigrated into a bisexual group. Within bi‐male groups, dominant males mated most frequently and males defended their groups during intergroup interactions. Lone females that visited groups or solitary males had a swelling more often than expected, and generally mated with the males they visited. Females had long receptive periods, several consecutive receptive cycles, and some overlap in receptive periods within groups. Females mated with extragroup males, and during infertile periods. We concluded that the males used the Diana monkeys for safety reasons and to obtain mating partners, and that female reproductive biology and behavior prevented the monopolization of groups of females by single males. Our data were inconclusive as regards the benefits to females of avoiding monopolization by males. Am. J. Primatol. 62:261–273, 2004. © 2004 Wiley‐Liss, Inc.     

Mating and oviposition behaviour in the legume pod borer, Maruca testulalis

Experiments were conducted outdoors and in the insect rearing laboratory at the International Institute of Tropical Agriculture (IITA) in Nigeria. Four or five nights pairing duration gave the highest mating percentage and oviposition by the pod borer. Direct observations on adult mating revealed that some males mated more than once. The majority of females mated only once. A one-to-one ratio (with 10 males: 10 females) gave the best results for mating and oviposition. Mating took place between 21.00 h and 05.00 h when temperature ranged between 20–25°C and rh over 80% with peak activity occurring between 02.00 and 03.00 h. Oviposition followed a similar trend to that observed for mating with an equal sex ratio (of 10 or more pairs per cage) giving the highest egg count. The average number of eggs laid was ca. 400 per female. These results show a much higher oviposition potential for Maruca testulalis than has been previously reported. The results are discussed with regard to the rearing of this species under laboratory conditions.

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Accouplement et comportement de ponte de Maruca testulalis

Résumé Tant de l'extérieur qu'au laboratoire (à l'IITA au Nigéria), la durée optimale de maintien des couples de M. testulalis Geyer (Lep. Pyralidae) pour obtenir l'accouplement et la ponte a été de 4 à 5 nuits, toutefois les meilleurs résultats ont été obtenus à l'extérieur. L'observation directe a révélé que contrairement aux femelles, les mâles peuvent copuler plus d'une fois. Le meilleur rapport sexuel pour obtenir l'accouplement et la ponte est 1/1 (10 mâles/10 femelles), suivi de près par un raport avec excédent de mâles. L'accouplement a lieu entre 21 h et 5 h quand la température est entre 20 et 30°C et HR supérieure à 80%; un pic est observé entre 2 et 3 h. La ponte présente la même tendance que l'accouplement, avec une production d'ufs plus élevée pour au moins 10 couples par cage. Le nombre moyen d'ufs par femelle est environ 400. Ces résultats sont supérieurs à ce qui avait été indiqué précédemment. La discussion porte sur les difficultés rencontrées pour élever cette espèce au laboratoire.

     

Genetic load,nutrient limitation,and seed production in Lupinus texensis (Fabaceae)

The interaction of mating system and nutrient limitation in determining seed production was investigated in the annual, self-compatible plant Lupinus texensis (Fabaceae). Abortion of developing seeds is a major factor limiting seed production in natural populations (17-28%). Selfing rates are generally low (0.02-0.21), suggesting that deleterious recessive genes may be maintained at significant levels in natural populations. The average inbreeding depression associated with seed development is δ = 0.24. Nutrient limitation reduced seed output across experimental treatments by a factor of 0.22 through decreased production of inflorescences, flowers, and ovules, and by a factor of 0.29 through increased abortion of fruits and of seeds within fruits. Competition for resources among fruits increased the frequency of seed abortion. Moreover, a greater proportion of selfed seeds were aborted as the overall abortion rate increased. Estimates of genetic load may therefore only be appropriate if undertaken in the field, and inbreeding depression may vary from year to year simply due to changes in environmental conditions rather than to underlying genetic changes in populations. The existence of inbreeding depression and the high frequency of abortions suggest that selective abortion favoring outcrossed progeny occurs in natural populations of L. texensis.     

Determinants of quaternary association in legume lectins

It is well known that the sequence of amino acids in proteins code for its tertiary structure. It is also known that there exists a relationship between sequence and the quaternary structure of proteins. The question addressed here is whether the nature of quaternary association can be predicted from the sequence, similar to the three-dimensional structure prediction from the sequence. The class of proteins called legume lectins is an interesting model system to investigate this problem, because they have very high sequence and tertiary structure homology, with diverse forms of quaternary association. Hence, we have used legume lectins as a probe in this paper to (1) gain novel insights about the relationship between sequence and quaternary structure; (2) identify the sequence motifs that are characteristic of a given type of quaternary association; and (3) predict the quaternary association from the sequence motif.     

Altitudinal variation in flowering area and position and their ecological significances of an alpine cushion Arenaria polytrichoides,a gynodioecious herb

Aims As foundation species in the alpine ecosystems, the reproduction and recruitment of alpine cushion plants are very important for sustaining the alpine ecosystem functions. However, it still remains unclear that how cushion plants effectively allocate resources to optimize reproductive fitness.Methods Here we selected five populations of a gynodioecious herb Arenaria polytrichoides with different exposures and slopes along an altitudinal gradient on the Baima snow mountain in northwest Yunnan, southwest China, to investigate and compare flowering area and positions, within and among populations and between female and hermaphroditic morphs. By doing so, we further discuss how the environmental stresses affect the cushion’s flowering attributes thus the population-level reproduction.Important findings The results showed that, individual plant size and resources allocated to flowering (flowering area %) both decreased with increasing elevation, indicating that the reproductive allocation strategy was significantly affected by elevation. However, a population at lower elevation showed lower reproductive investment than higher populations, suggesting that elevation was not the only factor affecting the cushion’s reproductive allocation. In addition, absolute flowering area increased with increasing individual size, but the flowering area ratio decreased, indicating that the increases in reproductive allocation are fewer than that in vegetative allocation. Hermaphroditic individuals invested more resources to flowering than females did, but again, such effect was affected by elevation. Moreover, within a single population, the flowering areas were significantly different among the four directions (east, south, west and north) within one single individual canopy, but such differences varied in different populations.     

Nodulation and N2 fixation byInga jinicuil,a woody legume in coffee plantations

Summary Nodule biomass and yearly C₂H₂ reduction rates are reported forInga jinicuil, a leguminous tree used for shade in Mexican coffee plantations. Annual fixation by this species approximates 35 kg ha^–1; which, when compared to nitrogen additions from fertilizers, represents an important nitrogen input to the coffee ecosystem.     

Variable selection in Platanthera bifolia (Orchidaceae): phenotypic selection differed between sex functions in a drought year

We estimated selection on three morphological characters in the hermaphroditic, hawkmoth-pollinated orchid Platanthera bifolia and explored selection surfaces through male and female function. The work was carried out in northern Sweden during two flowering seasons (1994 and 1995) in one natural population and one season (1995) in another natural population. Fitness was estimated as number of pollinia removed (male function) and number of fruits produced (female function). We detected directional selection towards larger inflorescence size (flower number) through both sex functions in both populations in 1995. In 1994, with an unusually dry growing season, 78% of the individuals failed to set any fruit, and there was selection for larger inflorescences only through male function. In this year, there was selection towards longer flower spurs, which could be a direct or indirect effect of spurs being shortened by drought. The results demonstrate that selection patterns may vary temporally and spatially, and that the 'male function hypothesis' may be applicable as female function is more resource dependent than male function.     

Breeding ecology of an Old World high-altitude warbler, Phylloscopus affinis

Reproduction of Tickell’s Leaf Warblers Phylloscopus affinis was studied in an alpine valley (29°27′N, 91°40′E, 3,980–5,600 m) in the Lhasa mountains, Tibet, at the upper elevational limit of the species’ breeding range. This species is a summer breeder, and is the only breeding Phylloscopus species in the valley. It nested in all types of shrubby vegetation across the altitudinal range of the valley. Most nests were placed close to the ground (<1 m) in low thorn bushes. Egg-laying dates fell between late May and early July, most within the first 3 weeks after the commencement of breeding. Mean clutch size was 4.0 (3–5) and mean brood size at fledging 3.4 (2–5). Incubation was by the females and lasted 13–14 days, and both parents cared for the young for 14–17 days. Nestlings ready to leave the nest were 13% heavier than the adults. Overall, 76% of nesting attempts produced at least one fledged young. Some aspects of the breeding biology of this high-altitude warbler were compared with those of lower-altitude Phylloscopus species.