Identification of marker-trait associations for self-compatibility in a segregating mapping population of Theobroma cacao L.

Increasing yield, quality, and disease resistance are important objectives for cacao breeding programs. Yield reduction in material improved for other traits is one of the main constraints caused by self-incompatibility (SI). Genes regulating SI in cacao have not been identified; therefore, knowledge of the location of genetic markers for and the effects of alleles determining SI will be useful for selecting uniformly self-compatible cultivars with higher yields. In a mapping population originating from a cross between a self-incompatible clone, Pound-7, and a self-compatible clone, UF-273, we observed important differences in flower retention at 15, 21, and 28 days after pollination. Our results suggest that the best time to measure flower retention is 15 days after pollination or later and that selecting self-compatibility (SC) thresholds is genotype specific. Using marker-trait association analysis we identified one marker, mTcCIR222, strongly associated with SC, as well as three surrounding markers (mTcCIR168, mTcCIR115 and mTcCIR158), all located near the proximal end of linkage group 4. These markers are currently being tested in our marker-assisted selection program.     

Loss of gametophytic self-incompatibility with evolution of inbreeding depression

Gametophytic self-incompatibility (SI) in plants is a widespread mechanism preventing self-fertilization and the ensuing inbreeding depression, but it often evolves to self-compatibility. We analyze genetic mechanisms for the breakdown of gametophytic SI, incorporating a dynamic model for the evolution of inbreeding depression allowing for partial purging of nearly recessive lethal mutations by selfing, and accounting for pollen limitation and sheltered load linked to the S-locus. We consider two mechanisms for the breakdown of gametophytic SI: a nonfunctional S-allele and an unlinked modifier locus that inactivates the S-locus. We show that, under a wide range of conditions, self-compatible alleles can invade a self-incompatible population. Conditions for invasion are always less stringent for a nonfunctional S-allele than for a modifier locus. The spread of self-compatible genotypes is favored by extremely high or low selfing rates, a small number of S-alleles, and pollen limitation. Observed parameter values suggest that the maintenance of gametophytic SI is caused by a combination of high inbreeding depression in self-incompatible populations coupled with intermediate selfing rates of the self-compatible genotypes and sheltered load linked to the S-locus.     

Evolution of floral scent in relation to self-incompatibility and capacity for autonomous self-pollination in the perennial herb Arabis alpina

Background and AimsThe transition from outcrossing to selfing is a frequent evolutionary shift in flowering plants and is predicted to result in reduced allocation to pollinator attraction if plants can self-pollinate autonomously. The evolution of selfing is associated with reduced visual floral signalling in many systems, but effects on floral scent have received less attention. We compared multiple populations of the arctic–alpine herb Arabis alpina (Brassicaceae), and asked whether the transition from self-incompatibility to self-compatibility has been associated with reduced visual and chemical floral signalling. We further examined whether floral signalling differ between self-compatible populations with low and high capacity for autonomous self-pollination, as would be expected if benefits of signalling decrease with reduced dependence on pollinators for pollen transfer.MethodsIn a common garden we documented flower size and floral scent emission rate and composition in eight self-compatible and nine self-incompatible A. alpina populations. These included self-compatible Scandinavian populations with high capacity for autonomous self-pollination, self-compatible populations with low capacity for autonomous self-pollination from France and Spain, and self-incompatible populations from Italy and Greece.Key ResultsThe self-compatible populations produced smaller and less scented flowers than the self-incompatible populations. However, flower size and scent emission rate did not differ between self-compatible populations with high and low capacity for autonomous self-pollination. Floral scent composition differed between self-compatible and self-incompatible populations, but also varied substantially among populations within the two categories.ConclusionsOur study demonstrates extensive variation in floral scent among populations of a geographically widespread species. Contrary to expectation, floral signalling did not differ between self-compatible populations with high and low capacity for autonomous self-pollination, indicating that dependence on pollinator attraction can only partly explain variation in floral signalling. Additional variation may reflect adaptation to other aspects of local environments, genetic drift, or a combination of these processes.     

Genome-wide association mapping of sexual incompatibility genes in cacao (<Emphasis Type="Italic">Theobroma cacao</Emphasis> L.)

Sexual compatibility limits the production of cacao plantations, being an important selection criterion in breeding programs. However, the current method for characterizing compatibility, based on the frequency of flower setting after controlled pollination, is time consuming, requiring a long time to identify self-compatible individuals. The identification of molecular markers in genomic regions can be an alternative to allow early selection of self-compatible plants. The present study aimed to identify SNP markers associated with sexual compatibility in cacao, by utilizing genome-wide association (GWAS) mapping. A population of 295 individuals mostly from third-generation breeding populations, but also founder clones, was used. This population was phenotypically characterized by hand pollinating 8199 flowers and evaluating the flower retention 15 days after pollination. In addition, leaf samples of each individual were collected and DNA extracted for genotyping by sequencing, generating 5301 SNP markers after cleaning. Genome-wide association mapping analysis was performed using Synbreed, GCTA, and TASSEL softwares. Significant markers associated to incompatibility, likely in strong linkage disequilibrium, were found within a region of 196 kb, in the proximal end of chromosome 4, suggesting the existence of a major gene in that region. However, this result should be validated in a larger population, considering that only 295 trees were used here. When the SNP effects were treated as random in the estimation process, many other regions in the genome appears to be involved with sexual incompatibility in cacao. Candidate genes were found not only in the proximal end of chromosome 4 but also spread in several other regions of the genome.     

山莨菪(茄科)的传粉生物学

茄科的多数种类具有自交不亲和的特点, 主要通过异花传粉结实; 但是, 一些物种或者物种内的部分种群或者个体却高度自交亲合, 转变为自交的繁育系统。该科植物山莨菪(Anisodus tanguticus)主要分布在青藏高原, 开花较早, 比其他晚开花的植物种类更加缺少有效的异花传粉昆虫。我们选择了位于不同海拔高度的2个种群进行比较研究, 主要目的是检验该物种的繁育系统是否在极端环境下由于传粉者的缺乏而发生了部分改变。研究发现,山莨菪的花不完全雌性先熟, 柱头和花药间的平均距离随着花开放时间的延长而不断缩小, 但两者在多数花的单花花期结束时并没有发生接触。因此, 山莨菪花主要表现为适应异花传粉的雌雄异位特征。然而, 少数花 (4.9％)的柱头和花药发生接触, 为“自动自交”的传粉解除了空间隔离。2个种群的多数个体存在自交不亲和机制, 应具有异花传粉的繁育系统; 但是部分个体具有明显的自交亲和能力, 为自交提供了生理基础。高海拔种群的传粉昆虫主要是厕蝇(Fannia sp.), 它们在不同植株间的活动能够保证异花传粉结实; 同时该种群的部分个体存在“自动自交”。低海拔种群的主要访花昆虫是蚂蚁, 它们在花内的活动导致花粉在同一朵花内传递, 而引起“协助自交”; 而异花传粉昆虫厕蝇的访花频率则较高海拔种群低。两个种群的结实均由于异花传粉者不足而受到传粉限制。因此两种不同类型的自交机制为该早期开花植物异花访花昆虫的不足提供了一定程度上的繁殖补偿。     

The status of cacao (Theobroma cacao,sterculiaceae) in the western hemisphere

There are, at the present time, effectively no long-range, ongoing programs in any tropical country of the western hemisphere dedicated to the improvement of cacao (Theobroma cacao, Sterculiaceae). While some effort is currently made to obtain new acquisitions of cacao cultivars exhibiting desirable characteristics and to maintain genepools of these trees, there are few data from field trials to prove and substantiate these qualities. In addition, there is a growing concern regarding the disparities between predicted yields of cacao trees through the use of “hybrid” seed and from actual production under field conditions. This has stimulated an awareness of the current inadequate understanding of the genetics of cacao and the lack of comprehension as to which cultivars, under distinct ecological conditions, are precocious, resistant to disease, or heavy bearing, or indeed demonstrate those traits vital to the success of farming programs adapted to today’s market conditions. This paper examines the events that have led to the current status of selection, development, and breeding of cacao. Alternative approaches are suggested.     

Bird diversity in cacao farms and forest fragments of western Panama

Theobroma cacao plantings, when managed under the shade of rainforest trees, provide habitat for many resident and migratory bird species. We compared the bird diversity and community structure in organic cacao farms and nearby forest fragments throughout mainland Bocas del Toro, Panama. We used this dataset to ask the following questions: (1) How do bird communities using cacao habitat compare to communities of nearby forest fragments? (2) To what extent do Northern migratory birds use shaded cacao farms, and do communities of resident birds shift their abundances in cacao farms seasonally? (3) Do small scale changes in shade management of cacao farms affect bird diversity? Using fixed radius point counts and additional observations, we recorded 234 landbird species, with 102 species that were observed in both cacao and forest fragments, 86 species that were only observed in cacao farms, and 46 species that were restricted to forest fragments. Cacao farms were rich in canopy and edge species such as tanagers, flycatchers and migratory warblers, but understory insectivores were nearly absent from cacao farms. We observed 27 migratory species, with 18 species in cacao farms only, two species in forest only, and seven species that occurred in both habitats. In cacao farms, the diversity of birds was significantly greater where there was less intensive management of the canopy shade trees. Shade tree species richness was most important for explaining variance in bird diversity. Our study shows that shaded cacao farms in western Panama provide habitat for a wide variety of resident and migratory bird species. Considering current land use trends in the region, we suggest that action must be taken to prevent conversion away from shaded cacao farms to land uses with lower biodiversity conservation value.     

The influence of emergent trees on rainfall distribution in a cacao agroforest (Sulawesi,Indonesia)

Emergent trees may have an influence on the volume and the spatial distribution of water input into agroforestry stands and may thus affect water availability for the main crops. Our goal was to analyze the influence of such trees on rainfall distribution in a cacao agroforest area in the rainforest margin zone of Central Sulawesi, Indonesia. The emergent trees studied belong to the species Bischofia javanica (Phyllanthaceae) and were 15 m high remnants from the natural forest. A set of 96 throughfall gauges was systematically distributed underneath canopies of cacao only, and underneath canopies of cacao plus emergent trees (cacao plus trees). From an earlier study we knew that stemflow can safely be estimated with less than 1% of gross precipitation (Pg).     

Small reductions in corolla size and pollen: ovule ratio,but no changes in flower shape in selfing populations of the North American <Emphasis Type="Italic">Arabidopsis lyrata</Emphasis>

The shift from outcrossing to selfing is often accompanied by striking changes in floral morphology towards a “selfing syndrome”, which is characterized by flowers with reduction in size, pollen: ovule (P/O) ratio, and herkogamy. This study aims to test whether such changes have occurred in the North American Arabidopsis lyrata, which is of particular interest because of the relatively recent transitions to selfing in this system. Flower size, flower shape, herkogamy levels, P/O ratio, and floral integration of six self-incompatible (outcrossing) and six self-compatible (selfing) populations of A. lyrata were measured in a common environment using conventional and geometric morphometrics methods. Although selfers had on average 9.2% smaller corollas, 8.4% longer pistils, and 21.5% lower P/O ratios than outcrossers, there were no differences in shape, floral integration, and herkogamy between outcrossing and selfing populations. Moreover, most variation in floral traits was explained by population genetic background rather than by mating system. We conclude that selfing populations in A. lyrata have not evolved a selfing syndrome.     

Dispersion models and sampling of cacao mirid bug Sahlbergella singularis (Hemiptera: Miridae) on Theobroma Cacao in southern Cameroon

The spatio-temporal distribution of Sahlbergella singularis Haglung, a major pest of cacao trees (Theobroma cacao) (Malvaceae), was studied for 2 yr in traditional cacao forest gardens in the humid forest area of southern Cameroon. The first objective was to analyze the dispersion of this insect on cacao trees. The second objective was to develop sampling plans based on fixed levels of precision for estimating S. singularis populations. The following models were used to analyze the data: Taylor's power law, Iwao's patchiness regression, the Nachman model, and the negative binomial distribution. Our results document that Taylor's power law was a better fit for the data than the Iwao and Nachman models. Taylor's b and Iwao's β were both significantly >1, indicating that S. singularis aggregated on specific trees. This result was further supported by the calculated common k of 1.75444. Iwao's α was significantly <0, indicating that the basic distribution component of S. singularis was the individual insect. Comparison of negative binomial (NBD) and Nachman models indicated that the NBD model was appropriate for studying S. singularis distribution. Optimal sample sizes for fixed precision levels of 0.10, 0.15, and 0.25 were estimated with Taylor's regression coefficients. Required sample sizes increased dramatically with increasing levels of precision. This is the first study on S. singularis dispersion in cacao plantations. Sampling plans, presented here, should be a tool for research on population dynamics and pest management decisions of mirid bugs on cacao.     

The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.     

Genetic population structure of cacao plantings within a young production area in Nicaragua

Significant cocoa production in the municipality of Waslala, Nicaragua, began in 1961. Since the 1980s, its economic importance to rural smallholders increased, and the region now contributes more than 50% of national cocoa bean production. This research aimed to assist local farmers to develop production of high-value cocoa based on optimal use of cacao biodiversity. Using microsatellite markers, the allelic composition and genetic structure of cacao was assessed from 44 representative plantings and two unmanaged trees. The population at Waslala consists of only three putative founder genotype spectra (lineages). Two (B and R) were introduced during the past 50 years and occur in >95% of all trees sampled, indicating high rates of outcrossing. Based on intermediate allelic diversity, there was large farm-to-farm multilocus genotypic variation. GIS analysis revealed unequal distribution of the genotype spectra, with R being frequent within a 2 km corridor along roads, and B at more remote sites with lower precipitation. The third lineage, Y, was detected in the two forest trees. For explaining the spatial stratification of the genotype spectra, both human intervention and a combination of management and selection driven by environmental conditions, appear responsible. Genotypes of individual trees were highly diverse across plantings, thus enabling selection for farm-specific qualities. On-farm populations can currently be most clearly recognized by the degree of the contribution of the three genotype spectra. Of two possible strategies for future development of cacao in Waslala, i.e. introducing more unrelated germplasm, or working with existing on-site diversity, the latter seems most appropriate. Superior genotypes could be selected by their specific composite genotype spectra as soon as associations with desired quality traits are established, and clonally multiplied. The two Y trees from the forest share a single multilocus genotype, possibly representing the Mayan, 'ancient Criollo' cacao.     

Cacao domestication I: the origin of the cacao cultivated by the Mayas

Criollo cacao (Theobroma cacao ssp. cacao) was cultivated by the Mayas over 1500 years ago. It has been suggested that Criollo cacao originated in Central America and that it evolved independently from the cacao populations in the Amazon basin. Cacao populations from the Amazon basin are included in the second morphogeographic group: Forastero, and assigned to T. cacao ssp. sphaerocarpum. To gain further insight into the origin and genetic basis of Criollo cacao from Central America, RFLP and microsatellite analyses were performed on a sample that avoided mixing pure Criollo individuals with individuals classified as Criollo but which might have been introgressed with Forastero genes. We distinguished these two types of individuals as Ancient and Modern Criollo. In contrast to previous studies, Ancient Criollo individuals formerly classified as 'wild', were found to form a closely related group together with Ancient Criollo individuals from South America. The Ancient Criollo trees were also closer to Colombian-Ecuadorian Forastero individuals than these Colombian-Ecuadorian trees were to other South American Forastero individuals. RFLP and microsatellite analyses revealed a high level of homozygosity and significantly low genetic diversity within the Ancient Criollo group. The results suggest that the Ancient Criollo individuals represent the original Criollo group. The results also implies that this group does not represent a separate subspecies and that it probably originated from a few individuals in South America that may have been spread by man within Central America.     

Microbiology of the root region of coconut and cacao under mixed cropping

Summary Mixed cropping of cacao in coconut plantations improved the microbial activity in the rhizosphere of coconut which may be attributed to an increase in organic matter content of soil due to periodic shedding of cacao leaves. When compared to coconut cultivation without cacao, mixed cropping of coconut with cacao appeared to stimulate the population of bacteria and fungi in the rhizosphere of coconut, including the nitrogen fixing and phosphate solubilising bacteria. The occurrence of indole acetic acid producing Escherichia sp. on the root surface of coconut and the appearance of gibberellin(like-substance) producing Aspergillus flavus and A. fumigatus in the rhizosphere were other salient observations made in a pioneering study on the microbiology of the root region of plantation crops under mixed cropping. re]19750702     

Tree species richness and density affect parasitoid diversity in cacao agroforestry

In Brazil, cacao is mostly planted beneath shade trees. The diversity of shade trees varies from monospecific to highly diverse canopies, characteristic of pristine Atlantic Forest. This study evaluates the relationships between family richness of Hymenoptera-Parasitica and Chrysidoidea, and tree species richness and density, the species richness of herbaceous understorey, and the area and age of the cacao agroforestry system. We sampled 16 cacao agroforestry systems, with canopy diversity ranging from one to 22 tree species per hectare, in three seasons: summer (March), winter (August) and spring (November). Parasitoids were sampled using eight Malaise-Townes traps per site. Tree species richness and density were enumerated within 1 ha at each site, and herbaceous plant species richness was calculated in eight 1 m² plots, within the hectare. The number of parasitoid families increased with tree species richness and density in spring and summer, but decreased in winter. Neither species richness of herbaceous plants nor area and age of the system affected parasitoid family richness. We suggest that the increase of parasitoid diversity with tree species richness and density in warmer seasons reflects increasing heterogeneity and availability of resources. The decrease in parasitoid family number with tree density in winter may be due to local impoverishment of resources, leading to parasitoid emigration to neighbouring forest remnants. This result implies that a higher diversity of shade trees will help to maintain high parasitoid levels and, in consequence, higher levels of natural enemies of cacao pests, particularly in the warmer seasons. This prediction is borne out in the experience of cacao producers. The proper management of shade tree diversity will play a vital role in maintaining the sustainability of cacao agroforestry production systems in the tropics and, concurrently, will maintain high biodiversity values in these locations.     

Breeding systems in a temperate mediterranean-type climate montane sclerophyllous forest in central Chile

The frequency of dioecy in the predominantly biotically-pollinated native flora of a temperate montane sclerophyllous forest in central Chile, 33^oS, is determined. Experimental crosses and other tests were performed on a taxonomically diverse set of annual herbs, perennial herbs and woody species to detect genetic self-incompatibility, spontaneous selfing capacity and obligate agamospermy.
The overall frequency of dioecy in the community is 9%.Dioecy is unequally represented among life-forms, increasing in frequency with greater longevity: 0% in annual herbs; 2% in perennial herbs; 17% in shrubs; 57% in trees. Thirty-eight percent of 37 hermaphrodite species proved to be genetically self-incompatible. Self-incompatibility, like dioecy, increases in frequency with longevity: 0% in annual herbs; 50% in perennial herbs; 80% in shrubs. The self-compatible species showed a wide range of breeding habits from facultative outcrossing to strong autogamy. However, in most self-compatible species hand self-pollination increased fruit and seed set over spontaneous selfing. Obligate agamospermy was not detected among hermaphrodite specieS. Absolute and relative fecundity were strongly correlated with the degree of compatibility. However, reduced fecundity in obligate outbreeding species could not be unequivocally attributed to pollinator limitation.
For equivalent life-forms, the montane sclerophyllous forest exhibits proportionately more obligate outbreeding species than some tropical forests and other temperate communities. The strong tendency for obligate outcrossing in the Chilean community is seen as a result of interplay between life-history constraints and the intense abiotic and biotic pressures that are characteristic of the mediterranean-type climate community.     

A meta-analysis of the relation between mating system,growth form and genotypic diversity in clonal plant species

Next to its well-described ecological advantages, clonal growth in plants may incur fitness costs, which are associated with the effects of typically large clonal individuals on the patterns of pollen dispersal. These fitness costs include increased selfing and inbreeding depression in self-compatible species, and reduced mate availability in self-incompatible species. Although fitness costs may affect mating system evolution, there is currently no strong evidence available that either self-compatibility or self-incompatibility is associated with clonality. One reason for this may be the variety in growth forms (from guerrilla to phalanx habits) within clonal species, and the fact that growth form may strongly affect mating patterns. We present the results of a formal meta-analysis of 72 published studies, aiming at reporting genotypic diversities across studies and at relating mating system with clonal growth form and genotypic diversity. We found lower genotypic diversities in clonal self-incompatible species compared to self-compatible species, suggesting that mate availability may indeed be reduced in clonal self- incompatible species. We also cannot confirm that mating system is associated with clonal growth form.     

Breeding systems in a secondary deciduous forest in Venezuela: The importance of life form, habitat, and pollination specificity

Breeding systems and mating systems of plants in a previously studied secondary deciduous forest were reanalysed in the context of new data. In this analysis, we increased the number of plant species (up to approximately 25% of the plant species in the community), included other life forms (23 annual and perennial species, plus habitat disturbance categories), and considered information about pollinator specificity. The frequencies of species with different sexual systems in a sample of 51 species were 82% hermaphrodite, 14% monoecious, and 4% dioecious. The frequencies of breeding systems in the sample of 49 hermaphroditic and monoecious species were 53% self-incompatible and 47% self-compatible. Self-compatible species included seven partially self-compatible, three self-compatible non-autogamous, and 13 self-compatible autogamous species. None of the species evaluated proved to be agamospermous. Fifty-five percent of the species tested were obligate outbreeders. The proportion of self-incompatible species was higher among trees and shrubs than among annual herbs. The proportion of self-compatible species for perennial herbs and lianas was not different. The association between annual herbs and autogamy was not strong: seven of 13 species were autogamous, five were partially self-compatible, and one was self-incompatible. The main characteristics or factors associated with breeding system were life cycle and successional stage. Short-lived species were mostly self-compatible, and xenogamy tended to be associated with forest and forest-border. In contrast, pollination specificity and life form were not consistently related to breeding system (self-compatibility or self-incompatibility) and mating system (xenogamy or autogamy).     

Oecophylla smaragdina ants provide pest control in Australian cacao

Ecosystem service‐driven methods of agricultural production that make optimal use of an ecosystem's natural services are often effective in smallholder crops, leading increases in habitat heterogeneity at large spatial scales that encourage beneficial species. Pollination services in cacao farms can be enhanced through efforts to conserve pollinator habitats, such as adding cacao fruit husks underneath trees to provide habitat and food resources for midge pollinators. However, it remains unclear how this habitat manipulation affects pest densities or biological control. Here, we evaluated the effects of the predaceous ant Oecophylla smaragdina on three major pests of Australian cacao (Helopeltis spp., Amblypelta spp., and Rhyparida nitida) in an Australian cacao farm, in the presence and absence of cacao fruit husks. We also evaluated the effect of O. smaragdina on their potentially plant‐damaging, mutualistic mealybug (Pseudococcidae) partners in the presence and absence of cacao fruit husks. Oecophylla smaragdina reduced the combined densities of Helopeltis spp. and Amblypelta spp., regardless of cacao fruit husk addition. The ants also reduced the densities of R. nitida in the absence of cacao fruit husks, but R. nitida densities were naturally low in the experimental plots that received the addition of cacao fruit husks, regardless of ant presence. In contrast, ants generally increased the densities of mealybugs. These data suggest that O. smaragdina ants provide pest control services for three major insect pests in Australian cacao farms. Furthermore, this control may be compatible with habitat manipulations (i.e., the addition of cacao fruit husks) to improve pollinator habitat.     

The genetics of self-incompatibility inLeavenworthia crassa rollins (Cruciferae)

Six plants of a self-incompatible population ofLeavenworthia çrassa were grown from seed collected in nature and cross-pollinated in all combinations. The incompatibility relationships between sibs were determined in eleven of the F₁ families. A one-locus sporophytic incompatibility system was established. None of the parents was homozygous at the S locus. At least five, and possibly all six, of the parents did not share an S allele. Only one pair of alleles was shown to have different interactions in the pollen and stigmata. The identity and expression of the S alleles were determined in six families. Eight pairs of alleles were independently expressed in both the pollen grains and the stigmata. Sixteen pairs of alleles showed dominance of one allele over the other in the pollen grains or the stigmata or both.F₁ plants of two crosses between different self-incompatible races were self-incompatible. F₁ plants of six crosses between self-incompatible and self-compatible races were self-incompatible; in five of the families, the frequency of pseudo-compatibility was higher than in the self-incompatible parent. Self-incompatible hybrids from a cross between a self-incompatible and a self-compatible population provide a method for rapidly determining allelic interactions in plants with a sporophytic incompatibility system.The research was carried out at the Biological Laboratories, Harvard University, Cambridge, Massachusetts, U.S.A.