Carbon-13 turnover in 'roxo de valinhos' fig trees

The aim of this study was to assess carbon-13 turnover in different organs of the fig tree, 'Roxo de Valinhos' cultivar. The experiment was carried out in an orchard at School of Agronomical Sciences, FCA/UNESP, Botucatu Campus, State of São Paulo, Brazil. The main photosynthetically active leaf was previously determined based on gas exchanges by means of an open portable photosynthesis system, IRGA. That leaf was placed in a chamber where the enriched gas injection occurred. The leaf enrichment time was 30 minutes. Treatments were constituted of seven fig trees removed from the soil after: 6; 24; 48; 72; 120; 168 and 360 hours of enrichment using (13)C, and their parts were sectioned into: apical bud, young leaves, adult leaves (photosynthetically active), lateral sprouts, fruits, and branch. The results allowed the establishment of the carbon-13 metabolism sequence in the studied parts: Young leaves > Fruits > Sprouts > Adult leaves > Apical bud > branch > Labeled leaf. 'Roxo de Valinhos' fig trees, had (13)C turnover of 24 hours and carbon-13 half-time shorter than 11 hours.

Effects of irrigation and nitrogen levels on qualitative and nutritional aspects of fig-trees (Ficus carica L.)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Water availability determines the richness and density of fig trees within Brazilian semideciduous forest landscapes

The success of fig trees in tropical ecosystems is evidenced by the great diversity (+750 species) and wide geographic distribution of the genus. We assessed the contribution of environmental variables on the species richness and density of fig trees in fragments of seasonal semideciduous forest (SSF) in Brazil. We assessed 20 forest fragments in three regions in Sao Paulo State, Brazil. Fig tree richness and density was estimated in rectangular plots, comprising 31.4 ha sampled. Both richness and fig tree density were linearly modeled as function of variables representing (1) fragment metrics, (2) forest structure, and (3) landscape metrics expressing water drainage in the fragments. Model selection was performed by comparing the AIC values (Akaike Information Criterion) and the relative weight of each model (wAIC). Both species richness and fig tree density were better explained by the water availability in the fragment (meter of streams/ha): wAICrichness = 0.45, wAICdensity = 0.96. The remaining variables related to anthropic perturbation and forest structure were of little weight in the models. The rainfall seasonality in SSF seems to select for both establishment strategies and morphological adaptations in the hemiepiphytic fig tree species. In the studied SSF, hemiepiphytes established at lower heights in their host trees than reported for fig trees in evergreen rainforests. Some hemiepiphytic fig species evolved superficial roots extending up to 100 m from their trunks, resulting in hectare-scale root zones that allow them to efficiently forage water and soil nutrients. The community of fig trees was robust to variation in forest structure and conservation level of SSF fragments, making this group of plants an important element for the functioning of seasonal tropical forests. © 2013 Elsevier Masson SAS. All rights reserved.

Memoranda respecting the culture of fig trees in the open air in England.

Transactions of the Horticultural society of London, 1820, iii, 74-85.

How to be a dioecious fig: Chemical mimicry between sexes matters only when both sexes flower synchronously

In nursery pollination mutualisms, which are usually obligate interactions, olfactory attraction of pollinators by floral volatile organic compounds (VOCs) is the main step in guaranteeing partner encounter. However, mechanisms ensuring the evolutionary stability of dioecious fig-pollinator mutualisms, in which female fig trees engage in pollination by deceit resulting in zero reproductive success of pollinators that visit them, are poorly understood. In dioecious figs, individuals of each sex should be selected to produce odours that their pollinating wasps cannot distinguish, especially since pollinators have usually only one choice of a nursery during their lifetime. To test the hypothesis of intersexual chemical mimicry, VOCs emitted by pollen-receptive figs of seven dioecious species were compared using headspace collection and gas chromatography-mass spectrometry analysis. First, fig-flower scents varied significantly among species, allowing host-species recognition. Second, in species in which male and female figs are synchronous, intersexual VOC variation was not significant. However, in species where figs of both sexes flower asynchronously, intersexual variation of VOCs was detectable. Finally, with one exception, there was no sexual dimorphism in scent quantity. We show that there are two ways to use scent to be a dioecious fig based on differences in flowering synchrony between the sexes.

Interference Competition and High Temperatures Reduce the Virulence of Fig Wasps and Stabilize a Fig-Wasp Mutualism

Fig trees are pollinated by fig wasps, which also oviposit in female flowers. The wasp larvae gall and eat developing seeds. Although fig trees benefit from allowing wasps to oviposit, because the wasp offspring disperse pollen, figs must prevent wasps fr

Fragmentation effects on diversity of wasp community and its impact on fig/fig wasp interaction in Ficus racemosa L.

Habitat fragmentation usually results in alteration of species composition or biological communities. However, little is known about the effect of habitat fragmentation on the fig/fig wasp system. In this study, we compared the structure of a fig wasp community and the interaction between figs and fig wasps of Ficus racemosa L. in a primary forest, a locally fragmented forest and a highly fragmented forest. Our results show that, in the highly fragmented forest, the proportion of pollinator wasps is lower and the proportion of non-pollinator wasps is higher compared with the primary forest and locally fragmented forest. The proportion of fruits without pollinator wasps in mature fruits is also greatly increased in the highly fragmented forest. The proportion of galls in all female flowers increases in the highly fragmented forest, whereas the proportion of viable seeds does not change considerably. The disruption of groups of fig trees results in a decrease in pollinator wasps and even might result in the extinction of pollinator wasps in some extreme cases, which may transform the reciprocal interaction between figs and fig wasps into a parasite/host system. Such an effect may lead to the local extinction of this keystone plant resource of rain forests in the process of evolution, and thereby, may change the structure and function of the tropical rain forest.

Speciation in fig wasps

There are over 700 species of fig trees in the tropics and several thousand species of fig wasps are associated with their syconia (inflorescences). These wasps comprise a monophyletic family of fig pollinators and several diverse lineages of non-pollinating wasps. The pollinator larvae gall fig flowers, while larvae of non-pollinating species either initiate their own galls or parasitise the galls of other wasps. A single fig species has 1-4 pollinator species and also hosts up to 30 non-pollinating wasp species. Most wasps show a high degree of host plant specificity and are known from only a single fig species. However, in some cases wasps may be shared across closely related fig species. There is impressive morphological coevolution between figs and fig wasps and this, combined with a high degree of partner specificity, led to the expectation that figs and pollinators have cospeciated extensively. Comparison of deep phylogenies supports long-term codivergence of figs and pollinators, but also suggests that some host shifts have occurred. Phylogenies of more closely related species do not match perfectly and may even be incongruent, suggesting significant roles for processes other than strict cospeciation. Combined with recent evidence on host specificity patterns, this suggests that pollinator wasps may often speciate by host shifts between closely related figs, or by duplication (the wasp speciates but the fig doesn't). The frequencies and biological details of these different modes of speciation invite further study. Far less is known about speciation in non-pollinating fig wasps. Some lineages have probably coevolved with figs and pollinators for most of the evolutionary history of the symbiosis, while others appear to be more recent colonisers. Many species appear to be highly host plant specific, but those that lay eggs through the fig wall without entering the syconium (the majority of species) may be subject to fewer constraints on host-shifting than pollinators. There is evidence for substantial host shifting in at least one gens, but also evidence for ecological speciation on the same host plant by niche shifts in other cases. Finally, recent work has begun to address the issue of “community phylogeny” and provided evidence for long-term co-divergence of multiple pollinating and non-pollinating wasp lineages with their host figs.

Interference competition and high temperature reduce the ‘virulence’ of fig wasps and contribute to stability of a fig-wasp mutualism

Fig trees are pollinated by fig wasps, which also oviposit in female flowers. The wasp larvae gall and eat developing seeds. Although fig trees benefit from allowing wasps to oviposit, because the wasp offspring disperse pollen, figs must prevent wasps from ovipositing in all flowers, or seed production would cease, and the mutualism would go extinct. In Ficus racemosa, we find that syconia (‘figs’) that have few foundresses (ovipositing wasps) are underexploited in the summer (few seeds, few galls, many empty ovules) and are overexploited in the winter (few seeds, many galls, few empty ovules). Conversely, syconia with many foundresses produce intermediate numbers of galls and seeds, regardless of season. We use experiments to explain these patterns, and thus, to explain how this mutualism is maintained. In the hot summer, wasps suffer short lifespans and therefore fail to oviposit in many flowers. In contrast, cooler temperatures in the winter permit longer wasp lifespans, which in turn allows most flowers to be exploited by the wasps. However, even in winter, only in syconia that happen to have few foundresses are most flowers turned into galls. In syconia with higher numbers of foundresses, interference competition reduces foundress lifespans, which reduces the proportion of flowers that are galled. We further show that syconia encourage the entry of multiple foundresses by delaying ostiole closure. Taken together, these factors allow fig trees to reduce galling in the wasp-benign winter and boost galling (and pollination) in the wasp-stressing summer. Interference competition has been shown to reduce virulence in pathogenic bacteria. Our results show that interference also maintains cooperation in a classic, cooperative symbiosis, thus linking theories of virulence and mutualism. More generally, our results reveal how frequency-dependent population regulation can occur in the fig-wasp mutualism, and how a host species can ‘set the rules of the game’ to ensure mutualistic behavior in its symbionts.

Longevity, early emergence and body size in a pollinating fig wasp - implications for stability in a fig-pollinator mutualism

1. Fig trees (Ficus) are pollinated only by agaonid wasps, whose larvae also gall fig ovules. Each ovule develops into either a seed (when pollinated) or a wasp (when an egg is also laid inside) but not both. 2. Ovipositing wasps (foundresses) favour ovules near the centre of the enclosed inflorescence (syconium or 'fig'), leaving ovules near the outer wall to develop into seeds. This spatial stratification of wasps and seeds ensures reproduction in both partners, and thereby enables mutualism persistence. However, the mechanism(s) responsible remain(s) unknown. 3. Theory shows that foundresses will search for increasingly rare inner ovules and ignore outer ovules, as long as ovipositing in outer ovules is sufficiently slow and/or if inner ovules confer greater fitness to wasps. The fig-pollinator mutualism can therefore be stabilized by strong time constraints on foundresses and by offspring fitness gradients over variation in ovule position. 4. Female fig wasps cannot leave their galls without male assistance. We found that females in outer ovules were unlikely to be released. Inner ovules thus have added value to foundresses, because their female offspring are more likely to mate and disperse. 5. For those offspring that did emerge, gall position (inner/outer) and body size did not influence the order in which female pollinators exited syconia, nor did early emerging wasps enjoy increased life spans. 6. We also found that the life spans of female wasps nearly doubled when given access to moisture. We suggest that conflict resolution in the fig-pollinator mutualism may thus be influenced by tropical seasonality, because wasps may be less able to over-exploit ovules in dry periods due to time constraints.

An extreme case of plant-insect co-diversification: figs and fig-pollinating wasps

It is thought that speciation in phytophagous insects is often due to colonization of novel host plants, because radiations of plant and insect lineages are typically asynchronous. Recent phylogenetic comparisons have supported this model of diversification for both insect herbivores and specialized pollinators. An exceptional case where contemporaneous plant insect diversification might be expected is the obligate mutualism between fig trees (Ficus species, Moraceae) and their pollinating wasps (Agaonidae, Hymenoptera). The ubiquity and ecological significance of this mutualism in tropical and subtropical ecosystems has long intrigued biologists, but the systematic challenge posed by >750 interacting species pairs has hindered progress toward understanding its evolutionary history. In particular, taxon sampling and analytical tools have been insufficient for large-scale co-phylogenetic analyses. Here, we sampled nearly 200 interacting pairs of fig and wasp species from across the globe. Two supermatrices were assembled: on average, wasps had sequences from 77% of six genes (5.6kb), figs had sequences from 60% of five genes (5.5 kb), and overall 850 new DNA sequences were generated for this study. We also developed a new analytical tool, Jane 2, for event-based phylogenetic reconciliation analysis of very large data sets. Separate Bayesian phylogenetic analyses for figs and fig wasps under relaxed molecular clock assumptions indicate Cretaceous diversification of crown groups and contemporaneous divergence for nearly half of all fig and pollinator lineages. Event-based co-phylogenetic analyses further support the co-diversification hypothesis. Biogeographic analyses indicate that the presentday distribution of fig and pollinator lineages is consistent with an Eurasian origin and subsequent dispersal, rather than with Gondwanan vicariance. Overall, our findings indicate that the fig-pollinator mutualism represents an extreme case among plant-insect interactions of coordinated dispersal and long-term co-diversification.

Biased oviposition and biased survival together help resolve a fig-wasp conflict

We report evidence that helps resolve two competing explanations for stability in the mutualism between Ficus racemosa fig trees and the Ceratosolen fusciceps wasps that pollinate them. The wasps lay eggs in the tree's ovules, with each wasp larva developing at the expense of a fig seed. Upon maturity, the female wasps collect pollen and disperse to a new tree, continuing the cycle. Fig fitness is increased by producing both seeds and female wasps, whereas short-term wasp fitness increases only with more wasps, thereby resulting in a conflict of interests. We show experimentally that wasps exploit the inner layers of ovules first (the biased oviposition explanation), which is consistent with optimal-foraging theory. As oviposition increases, seeds in the middle layer are replaced on a one-to-one basis by pollinator offspring, which is also consistent with biased oviposition. Finally, in the outer layer of ovules, seeds disappear but are only partially replaced by pollinator offspring, which suggests high wasp mortality (the biased survival or ‘unbeatable seeds’ explanation). Our results therefore suggest that both biased oviposition and biased survival ensure seed production, thereby stabilizing the mutualism. We further argue that biased oviposition can maintain biased survival by selecting against wasp traits to overcome fig defenses. Finally, we report evidence suggesting that F. racemosa balances seed and wasp production at the level of the tree. Because figs are probably selected to allocate equally to male and female function, a 1:1 seed:wasp ratio suggests that fig trees are in control of the mutualism.

How to be a fig wasp down under: the diversity and structure of an Australian fig wasp community

Endophytic insects and their parasitoids provide valuable models for community ecology. The wasp communities in inflorescences of fig trees have great potential for comparative studies, but we must first describe individual communities. Here, we add to the few detailed studies of such communities by describing the one associated with Ficus rubiginosa in Australia. First, we describe community composition, using two different sampling procedures. Overall, we identified 14 species of non-pollinating fig wasp (NPFW) that fall into two size classes. Small wasps, including pollinators, gallers and their parasitoids, were more abundant than large wasps (both galler and parasitoid species). We show that in figs where wasps emerge naturally, the presence of large wasps may partly explain the low emergence of small wasps. During fig development, large gallers oviposit first, before and around the time of pollination, while parasitoids lay eggs after pollination. We further show that parasitoids in the subfamily Sycoryctinae, which comprise the majority of all individual NPFWs, segregate temporally by laying eggs at different stages of fig development. We discuss our results in terms of species co-existence and community structure and compare our findings to those from fig wasp communities on other continents.

Genetic characterization of fig tree mutants with molecular markers

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Fig tree orchard formation using organic fertilization

The present work aimed at evaluating the effects of cattle manure fertilization on the growth and yield of fig trees. The cultivar 'Roxo de Valinhos' was used. One-, two-, three-and four-year-old trees were treated with cattle manure containing 0, 25, 50, 75, 100, 125 and 150% of the recommended nitrogen level per plant. The experiment was carried out using randomized blocks with 7 treatments, 5 replicates and 5 plants per experimental plot. The evaluated characteristics were: plant height, stem diameter, secondary branch length and diameter, yield per plant (kg plant-1), besides some quality characteristics of fruits such as pH, titratable acidity, soluble solids and texture. Manure application enhanced plant growth and fruit production. Significant differences were observed only for soluble solids content, pH and texture, which varied according to the crop cycle. After four crop cycles (2002/03, 2003/04, 2004/05 and 2005/06), the best results (about 5.0 kg of fruits per plant) were obtained with 100% of the recommended nitrogen dose, which corresponded to 14.3 kg of cattle manure per plant, in the last crop cycle (2005/2006).