Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.     

The effect of unripe fruits on ripe fruit removal by birds in Pistacia terebinthus: flag or handicap?

The shrub Pistacia terebinthus produces crowded infructescences with up to several hundred fruits, which are bright red when unripe and turn green when ripe. Most fruits contain an empty seed and never reach maturity. More ripe fruits were removed by birds from experimental bicolored fruit displays (consisting of infructescences with ten ripe fruits and stripped of unripe fruits, paired with infructescences with only unripe fruits) than from monocolored ones (single infructescences with ten ripe fruits and stripped of unripe fruits). Thus, the presence of unripe fruits seems to increase the conspicuousness or attractiveness of fruit displays to fruit-eating birds. A second experiment compared ripe fruit removal from experimental infructescences having only ripe fruits, with that from control infructescences containing both ripe fruits and natural numbers of unripe fruits, all on P. terebinthus plants. Unlike the first experiment, each bicolored display in this case consisted of a single infructescence with both unripe and ripe fruits. A higher proportion of ripe fruits was removed by birds from infructescences free of unripe fruits. This result suggests that the presence of unripe fruits reduces the accessibility of ripe fruits for fruit-eating birds. This is further supported by field observations of bird foraging behavior.     

Maturation characteristics of Rubus pennsylvanicus fruit: are black and red the same?

Summary Fruit of the blackberry, Rubus pennsylvanicus Poir. (Rosaceae), were examined to determine variation in maturation characteristics. Maturation timing and rate varied greatly among individual fruits, resulting in a bi-colored fruiting display comprised largely of two maturation stages, pre-ripe (salmon and scarlet) and ripe (dark brown and black). While ripe fruit were generally larger and heavier than pre-ripe fruit, exhibiting greater fresh and dry fruit weight, diameter, water content, and total seed weight, no significant differences were found in energy content, i.e. numbers of calories per gram pulp, or in pulp:seed ratio. The differences between ripe and pre-ripe fruit appear to be due largely to an increase in water content and seed weight with maturity. The fact that little energetic benefit accrues to the preferential selection of ripe fruit suggests that bi-colored Rubus displays may be considered to be unicolored.     

Fruit characteristics and factors affecting fruit removal in a Panamanian community of strangler figs

We describe fruiting characteristics for 12 species in a community of strangler figs (Moraceae: Urostigma) studied in Panama. We quantify diurnal and nocturnal removal rates and proportions of fruits removed, and relate them to the activities of the main dispersers of the figs: bats and birds. These results combined with previous studies show that there are clear differences between fig species with fruit that ripen red and those with fruit that remain green(ish). In the red-fruited species, the fruit are small, ripen asynchronously over relatively long periods, produce little scent, and are mainly taken during the day by birds. In contrast, in the green(ish)-fruited species, the fruits are larger, span a range of sizes, ripen relatively synchronously, produce very distinctive aromas, and are mainly taken at night by bats. This dichotomy in fruiting characteristics suggests coadaptive links between groups of dispersers and different species within the genus Ficus. All fig species produce a range of fruit crop sizes (10–155 fuits/m² canopy area) of which a high proportion were removed by seed dispersers (>80%). Removal rates (fruit removed per day) were positively correlated with crop size, suggesting that trees with large crop size attract more frugivores. Removal rates of green-fruited figs were significantly lower and persistence and abortion of ripe fruit were significant higher around full moon, apparently due to the reduced activity of bats. We further estimate the number of bats that are sustained by a tree fruit crop and account for the observed fruit removal. We then discuss the evidence for coadaptation between different groups of figs and their seed dispersers, Finally, we consider the conservation implications for figs as keystone resources in tropical forests. Received: 26 April 1999 / Accepted: 10 January 2000     

A field test of the directed deterrence hypothesis in two species of wild chili

The directed deterrence hypothesis posits that secondary metabolites in ripe fruit function to deter fruit consumption by vertebrates that do not disperse seeds, while not impacting consumption by those that do. We tested this hypothesis in two species of wild chilies (Capsicum spp.). Both produce fruits that contain capsaicinoids, the compounds responsible for the pungency of chilies. Previous work suggests seed-dispersing birds but not seed-destroying rodents consume chili fruits, presumably because rodents are deterred by capsaicin. However, fruit removal from chili plants by rodents and other mammals has not been previously explored. Because laboratory rodents can develop a preference for capsaicin, it is quite possible that wild rodents are natural consumers of chili fruits. We monitored the fate of 125 marked fruits of Capsicum chacoense and 291 fruits of Capsicum annuum. For both species, essentially all fruit removal occurred during the day, when rodents are inactive. Video monitoring revealed fruit removal only by birds, mostly by species known to disperse chili seeds in viable condition. Furthermore, these species are from taxonomic groups that tend to specialize on lipid-rich fruits. Both species of chili produce fruits that are unusually high in lipids (35% in C. chacoense, 24% in C. annuum). These results support the directed deterrence hypothesis and suggest that fruiting plants distinguish between seed predators and seed dispersers by producing fruits that repel the former and attract the latter.     

Does attraction to frugivores or defense against pathogens shape fruit pulp composition?

Fruit traits evolve in response to an evolutionary triad between plants, seed dispersers, and antagonists that consume fruits but do not disperse seeds. The defense trade-off hypothesis predicts that the composition of nutrients and of secondary compounds in fruit pulp is shaped by a trade-off between defense against antagonists and attraction to seed dispersers. The removal rate model of this hypothesis predicts a negative relationship between nutrients and secondary compounds, whereas the toxin-titration model predicts a positive relationship. To test these alternative models, we evaluated whether the contents of nutrients and secondary compounds can be used to predict fruit removal by mutualists and pathogens in 14 bird-dispersed plants on a subtropical island in São Paulo state, southeastern Brazil. We selected eight to ten individuals of each species and prevented fruit removal by covering four branches with a net and left fruits on four other branches available to both, vertebrate fruit consumers and pathogens. The persistence of ripe fruits was drastically different among species for bagged and open fruits, and all fruit species persisted longer when protected against seed dispersers. We found that those fruits that are quickly removed by vertebrates are nutrient-rich, but although the attack rate of pathogens is also high, these fruits have low contents of quantitative defenses such as tannins and phenols. Thus, we suggest that the fruit removal rate by seed dispersers is the primary factor selecting the levels of fruit defense. Likewise, nutrient-poor fruits have low removal of seed dispersers and low probability of attack by pathogens. These species retain ripe fruits in an intact condition for a prolonged period because they are highly defended by secondary compounds, which reduce overall attractiveness. However, this strategy might be advantageous for plants that depend on rare or unreliable dispersers.     

Does individual variation in fruit profitability override color differences in avian choice of red or white <Emphasis Type="Italic">Ilex serrata</Emphasis> fruits?

Although avian color preferences have been studied and documented in controlled experiments, they have not been demonstrated under natural conditions in most cases. We hypothesized that avian fruit choice reflects intraspecific variation in fruit characteristics other than color, rather than fruit color differences. By planting one Ilex serrata Thunb. (red form) and one I. serrata forma leucocarpa Beissner (white form), which produce red and white fruits, respectively, at each of five points, we examined the proportion of fruits removed per tree and fruit choice by three avian species based on fruit color and other fruit characteristics. The proportion of fruits removed increased with pulpy sugar concentration and fruit diameter, but it did not differ between fruit colors. The main foragers, resident brown-eared bulbuls Hypsypetes amaurotis, consumed fruits regardless of color, but correspondingly to fruit removal, and appeared to base their fruit choice on pulpy sugar concentration and fruit diameter rather than on color. In contrast, the minor foragers, migrant Daurian redstarts Phoenicurus auroreus (Pallas) and Siberian bluechats Tarsiger cyanurus (Pallas), tended to choose red fruits and were possibly attracted by them. In conclusion, fruit removal per tree reflected individual variation in fruit profitability more strongly than differences in fruit color, even though the individual variation was not remarkable. The importance of color in fruit choice differed based on species, residency status, and major/minor foragers.     

Factors affecting seed rain beneath fleshy-fruited plants

We investigated factors affecting seed rain beneath nine fleshy-fruited fruiting plant species growing in a 1-ha plot of planted Pinus thunbergii in central Japan. We tested whether the numbers of seeds and seed species dropped by birds beneath fruiting plants were correlated with the number of fruits removed by birds from the plants. Most of fruiting plant species with high fruit removal had significantly high seed rain. Both the numbers of seeds and seed species dropped were significantly, positively correlated with the number of fruits removed across for all fruiting plant species. Therefore, fruit removal predicted the difference among heterospecific fruiting plants in seed rain. We also tested whether the number of fruits removed from fruiting plants by birds was related with fruit crop size, fruit size, and height of the plants, and the numbers of fruits and fruit species of neighboring plants near the plants. Most of fruiting plant species with high fruit crop size had significantly high fruit removal. The number of fruits removed was significantly, positively correlated with both the fruit crop size and the number of neighboring fruits across the nine fruiting plant species. However, the effect of the neighboring fruit density on fruit removal was lower remarkably than that of fruit crop size. Therefore, fruit crop size best predicted the differences among heterospecific fruiting plants in fruit removal. We suggest that fruiting plant species with high fruit crop size and high fruit removal contribute to intensive seed rain beneath them. This revised version was published online in June 2006 with corrections to the Cover Date.     

Fruit removal of Coprosma quadrifida (Rubiaceae) by birds in south-eastern Australia

Abstract Removal of the fleshy fruits of Coprosma quadrifida by birds was examined in relation to fruit crop size, neighbouring fruit crop size and time in the season in montane wet sclerophyll forest in south-eastern Australia. On average, 84% of individual fruit crops were removed. Frugivores removed fruit in proportion to its abundance, indicating that they continually assess availability rather than respond to a critical abundance. The size of neighbouring fruit crops did not influence fruit removal rates, suggesting that there is little competition among neighbours for dispersers. The ripe fruits of C. quadrifida were eaten by 14 bird species, comprising 50% of the bird species trapped. Silvereyes (Zosterops lateralis) were the most abundant consumers of fruits. With the exception of the parrot, Platycercus elegans, all birds defecated seeds intact and had no influence on seed viability.     

Fruit removal efficiency and success: influence of crop size in a neotropical treelet

The reproductive success of animal-dispersed plants is closely linked to the number of seeds that they are able to disperse. The fruit crop size hypothesis states that a plant with large fruit crop size will attract more dispersers than a plant with a smaller fruit crop, which may result in more seeds being dispersed from the foremost. In this study, we experimentally examined the effect of crop size and other factors on primary seed dispersal in a neotropical shrub/tree, Casearia corymbosa (Flacourtiaceae). We used two predictive variables of reproductive success, which produce an accurate picture of seed dispersal ratio: fruit removal efficiency (proportion of a fruit crop removed by frugivores) and fruit removal success (relative contribution of each individual tree to the number of fruits removed in the population). We established two levels of fruit crop size at the C. corymbosa individuals, using plants with large (150 fruits) and small crops (50 fruits). We found that individual plants with larger crops had significantly higher values of fruit removal efficiency (92.6%) and success (5%) than plants with smaller crops (69.3% and 1.3%, respectively). Fruit removal efficiency was related to vegetation type, plant height and fruit width, but the variance explained by these variables was low ( < 8%). Fruit removal success was significantly related to crop size ( > 90% of the variance explained). These results suggest that fruit removal efficiency and success are strongly related to fruit crop size of C. corymbosa plants.     

Influence of between-year variation in the density of Rhus <Emphasis Type="Italic">trichocarpa</Emphasis> fruits on the removal of fruit by birds

The fruiting phenology and fruit removal patterns of Rhus trichocarpa Miq. (Anacardiaceae) were investigated in a warm-temperate secondary forest in Japan. Mature fruits of this species are dispersed by birds. Effects of fruit display size and canopy openness on fruit removal were investigated in years with different fruit densities (i.e., masting and non-masting years). Moreover, effects of increased canopy openness during winter on fruit removal were also investigated. Seasonal patterns of fruit removal were quite different between masting and non- masting years. In the non-mast year, fruits were removed by birds soon after maturation in the summer. In contrast, in the mast year, fruits were removed gradually by birds from summer to winter. Moreover, the rate of fruit removal was greater for trees with a larger display size in the non-mast year, whereas that was greater for trees with greater canopy openness in the mast year. Canopy openness increased in winter, and fruit removal in winter was enhanced in trees with a more open canopy only in the non-mast year. These observations strongly suggest that avian dispersers became satiated in the mast year, whereas fruit removal was enhanced in the non-mast year. In the mast year, although many fruits were not dispersed until winter, they were neither depredated nor rotten, and the long period of fruit removal by birds may have enhanced fruit dispersal. The large abundance of seedlings and saplings of this species in Japanese secondary forests suggests that this fruiting strategy is beneficial.     

Phenology of vertebrate removal ofMyoporum accuminatum fruits in a Queensland forest

Summary Fruit removal rates ofMyoporum acuminatum in a Queensland population were correlated with size of the ripe fruit crop throughout the season, but only sporadically correlated with nutritive contents of the fruits. Removal rates, as a proportion of crop size, varied synchronously among individuals and did not differ consistently among individuals with differing fruit traits. The principal avian consumer wasMeliphaga lewinii.     

Fruit size in a tropical tree species: variation,preference by birds,and heritability

Ocotea tenera (Lauraceae), an understory bird-dispersed tree, produces single-seeded fruits that vary in diameter from 1.4 to 2.4 cm. Much of the variation within a population at Monteverde, Costa Rica occurred within individual trees. The relative size of fruits produced by different trees remained generally constant over an 11-year period despite slight differences between years in the average size of fruits produced by a given tree.Fruit-eating birds could thus express their preferences for particular fruit size characteristics by choosing among trees that have distinct distributions of fruit diameters, and between individually variable fruits within trees. In a field study of individually marked fruits, birds removed 46.2% of fruits; the rest of the fruits were destroyed by invertebrate (25.3%) and vertebrate (4.3%) pulp-feeders or aborted by the plant after remaining ripe but uneaten for as long as 100 days (24.2%). The four major avian seed dispersers of O. tenera each have gape widths exceeding all but the largest fruits. Birds preferred plants with greater-than-average-sized fruits; within trees, they favored larger fruits, apparently because net pulp mass increases with fruit diameter. Fruits that ripened early in the season were more likely to be removed and were removed more quickly than late-ripening fruits.Based on mother-offspring regressions of mean fruit size, the phenotypic variation in fruit diameter in O. tenera is highly heritable, indicating the potential for an evolutionary response to selection by birds. Nonetheless, directional selection on fruit size or shape is likely to be inconsistent, constrained by genetic correlations, and weak compared to selection on traits like fecundity or phenology.     

A <Emphasis Type="Italic">Colletotrichum gloeosporioides</Emphasis>-induced esterase gene of nonclimacteric pepper (<Emphasis Type="Italic">Capsicum annuum</Emphasis>) fruit during ripening plays a role in resistance against fungal infection

Ripe fruits of pepper (Capsicum annuum) are resistant to the anthracnose fungus, Colletotrichum gloeosporioides, whereas unripe-mature fruits are susceptible. A pepper esterase gene (PepEST) that is highly expressed during an incompatible interaction between the ripe fruit of pepper and C. gloeosporioides was previously cloned. Deduced amino acid sequence of PepEST cDNA showed homology to both esterases and lipases, and contained -HGGGF- and -GXSXG- motifs and a catalytic triad. Inhibition of PepEST activity by a specific inhibitor of serine hydrolase demonstrated that a serine residue is critical for the enzyme activity. Expression of PepEST gene was fruit-specific in response to C. gloeosporioides inoculation, and up-regulated by wounding or jasmonic acid treatment during ripening. PepEST mRNA and protein was differentially accumulated in ripe vs. unripe fruit from 24 h after inoculation when C. gloeosporioides isinvading into fruits. Immunochemical examination revealed that PepEST accumulation was localized inepidermal and cortical cell layers in infected ripe fruit, but rarely even in epidermal cells in infected unripe one. Over-expression of PepEST in transgenic Arabidopsis plants caused restriction of Alternaria brassicicola colonization by inhibition of spore production, resulting in enhanced resistance against A.brassicicola. These results suggest that PepEST is involved in the resistance of ripe fruit against C.gloeosporioides infection.^†These authors contributed equally to the work     

The fruits of selectivity: how birds forage on Goupia glabra fruits of different ripeness

Although many studies have been published on avian fruit selection, few have addressed the effects of fruit scarcity on the patterns of fruit choice. Here, we compared the consumption of seven bird species for six simultaneously present maturation stages of Goupia glabra fruits. Ripe G. glabra fruits contain more lipids, carbohydrates and energy, and fewer phenols, than unripe fruits. All bird species selected from among ripening stages and removed a higher proportion of ripe fruits than of intermediate or unripe fruits. Importantly, however, fruit choice was flexible in all species. Whether birds preferred or avoided fruits of intermediate ripeness depended on the overall fruit supply. When ripe fruits were scarce, birds showed a higher acceptance of fruits of intermediate ripeness, but still rejected the least ripe fruit stages. In a foraging bout, most birds fed on fruits of the same ripeness. By doing so, birds maximised instantaneous energy gain per time, because search time was longer for riper fruits while energy intake was lower for less ripe fruits. The results suggest that birds select fruits based on fine-scale differences in profitability, but accept less profitable fruits during low fruit abundance. If environmental factors such as overall fruit availability influence avian fruit choice, we suggest that the potential for directional selective pressures on fruit compounds is restricted.     

Impact on fruit removal and seed predation of a secondary metabolite, emodin, in Rhamnus alaternus fruit pulp

There are two contradictory approaches to explaining the presence of secondary metabolites in ripe fruits. One holds that they evolved toward enhancing dispersal success (adaptive approach); the other claims that they evolved primarily to deter herbivores from eating leaves and seeds and that their presence in ripe fruits is a byproduct of that function (non‐adaptive approach). We tested the validity of three hypotheses of the adaptive approach that explain the presence of secondary metabolites in ripe fruits. We explored the current function of a secondary metabolite, emodin, in Mediterranean buckthorn (Rhamnus alaternus, Rhamnaceae) fruits by relating intraspecific variation and seasonal patterns of concentration to fruit removal and seed damage and by conducting feeding trials with captive birds presented artificial fruits that varied in emodin concentration. The concentration of emodin in the pulp of 10–13 Rhamnus plants from the same population was determined by HPLC every month during two fruiting seasons. Fruit removal by birds and seed predation by invertebrates and microbes were determined for the same plants. Emodin concentration rose during the first stages of ripening, reaching a peak before the fruits were ripe, and then decreased to a minimum when the fruits were ripe. No significant correlation between emodin concentration and ripe fruit removal rate among trees was observed in the first year, whereas in the second year the correlation was positive and significant. Thus, the impact of emodin on fruit selection varied between years, suggesting that emodin concentration does not solely govern fruit selection. A significant negative correlation was found in the first year between emodin concentration and seed predation during the first fruiting month. The yellow‐vented bulbul (Pycnonotus xanthopygos), a seed dispersing bird, distinguished between artificial foods that differed in emodin concentration (control, 0.001% and 0.002%), always preferring the lower concentration. In contrast, house sparrows, (Passer domesticus), a seed predator, did not detect such differences in emodin concentration but did distinguish between control foods and food with 0.005% and 0.001% emodin. We suggest that emodin has an ecological role, preventing seed predation by invertebrates and microbes without decreasing fruit removal by avian dispersers.     

An evaluation of vertebrate seed dispersal syndromes in four species of black nightshade (Solanum sect. Solanum)

We examined the ecological relevance of bird versus mammal dispersal syndromes in four species of Solanum, S. americanum Type A, S. americanum Type B, S. ptychanthum, and S. sarrachoides. These plants were selected because their morphological characteristics, such as fruit color, mass, and persistence, resembled those typically associated with classically-defined bird and mammal dispersal syndromes. We monitored persistence of tagged fruits, compared physical and chemical chaacteristics, performed fruit preference trials with northern bobwhite quail (Colinus virginianus), deer mice (Peromyscus maniculatus), and American robins (Turdus migratorius), and assessed differences in use of olfactory cues by foraging deer mice. We predicted that principal components analysis of physical and chemical characteristics would place fruits of these species along a clear gradient from bird to mammal dispersal syndromes (S. americanum Type A, S. americanum Type B, S. ptychanthum, and S. sarrachoides). However, physical and chemical characteristics did not consistently follow the gradient. Also, contrary to expectations, both birds and mammals demonstrated a preference for S. americanum Types A and B, both bird fruits. Deer mice and bobwhite quail showed much less discrimination among fruit types than did American robins. While the relatively strong odor of the green-fruited S. sarrachoides suggested a mammalian attractant, deer mice discovered the relatively odorless S. americanum Type A significantly more quickly. We conclude that in Solanum, suites of morphological characteristics resembling bird and mammal dispersal syndromes are not good predictors of fruit choice by birds and mammals. We note, however, that this conclusion is based on a sample of three animal species. Alternative explanations for fruit character suites should be considered. For example, the adaptive significance for an association of green coloration of ripe fruit with impersistence (dropping when ripe), as in S. sarrachoides and some S. ptychanthum, may relate more to photosynthesis and carbon balance in detached fruits than to disperser attraction.     

Ant seed removal in a non-myrmecochorous Neotropical shrub: Implications for seed dispersal

This study investigated ant seed removal of Piper sancti-felicis, an early successional Neotropical shrub. Neotropical Piper are a classic example of bat-dispersed plants, but we suggest that ants are underappreciated dispersal agents. We identified eleven ant species from the genera Aphaenogaster, Ectatomma, Paratrechina, Pheidole, Trachymyrmex, and Wasmannia recruiting to and harvesting P. sancti-felicis seeds in forest edge and secondary forest sites at La Selva, Costa Rica. We also tested for differences in ant recruitment to five states in which ants can commonly encounter seeds: unripe fruit, ripe fruit, overripe fruit, bat feces, and cleaned seeds. Overall, ants harvested more seeds from ripe and overripe fruits than other states, but this varied among species. To better understand the mechanisms behind ant preferences for ripe/overripe fruit, we also studied how alkenylphenols, secondary metabolites found in high concentrations in P. sancti-felicis fruits, affected foraging behavior in one genus of potential ant dispersers, Ectatomma. We found no effects of alkenylphenols on recruitment of Ectatomma to fruits, and thus, these compounds are unlikely to explain differences in ant recruitment among fruits of different maturity. Considering that P. sancti-felicis seeds have no apparent adaptations for ant dispersal, and few ants removed seeds that were cleaned of pulp, we hypothesize that most ants are harvesting its seeds for the nutritional rewards in the attached pulp. This study emphasizes the importance of ants as important additional dispersers of P. sancti-felicis and suggests that other non-myrmecochorous, vertebrate-dispersed plants may similarly benefit from the recruitment to fruit by ants.     

Petals may act as a reward: myrmecochory in shrubby Darwinia species of south‐eastern Australia

The removal rates of fruits (the diaspore) were assessed in exclusion experiments in three shrubby species of Darwinia, endemic to the Sydney Basin of south‐eastern Australia. There was a clear pattern of fruit removal in just a few days by small species of ants from the genera Iridomyrmex, Crematogaster and Pheidole. When only ants had access to fruits an average of 41% of fruits were removed, whereas when only vertebrates had access an average of 13% of fruits were removed. There was some variation in the number of fruits removed both between species and within species across study sites or annual seed crops. When ants had access to seeds this variation in removal amounted to 10–80% over 1–4 days. Uniquely in Darwinia, it is the old petals (which remain attached to the fruit) that are attractive to the ants, rather than a specific lipid body. This likely represents a case of successful ecological fitting by Darwinia to take advantage of the widespread presence of myrmecochory in the Australian flora.     

Altered Tomato (<Emphasis Type="Italic">Lycopersicon esculentum</Emphasis> Mill.) Fruit Cuticle Biomechanics of a Pleiotropic Non Ripening Mutant

Tomato (Lycopersicon esculentum Mill.) fruit ripening involves multiple metabolic changes resulting in softening and pigmentation. We investigated the mechanics and morphology of the enzymatically isolated cuticular membrane (CM) of cv. Ailsa Craig wild-type (wt) and nonripening mutant (nor) at three developmental stages. Cuticle thickness and degree of cutinization increased significantly from immature to fully ripe fruits for both wt and nor without differences between them. Mechanical characterization was carried out on dry and fully hydrated samples in uni-axial tension to determine their modulus of elasticity, stress, and strain at failure. Corresponding stress-strain diagrams were biphasic and showed yield for virtually all dry CM samples, while that of hydrated CM displayed considerable differences between wt and nor fruits. Concerning the mechanical properties, the CM of wt fruits was characterized by increasing stiffness and strength during fruit growth and maturation in both dry and hydrated states, whereas the CM of nor fruits was significantly less stiff and weaker at full maturity. Hydration generally caused lower moduli of elasticity and strength, while breaking strain was significantly affected only for the CM of ripe nor fruits. This plasticizing effect of water increased towards full maturity for both wt and nor, and may be related to fiber content in the CM matrix and hydration state of the cuticle. Comparative analysis of two additional wild-type tomato cultivars supported the ripening-related stiffening of the CM of Ailsa Craig wt and the altered mechanical properties of the nor mutant, as well as the plasticizing effect of water.