Preformation and distribution of staminate and pistillate flowers in growth units of Nothofagus alpina and N. obliqua (Nothofagaceae)

Background and Aims

The distribution and differentiation times of flowers in monoecious wind-pollinated plants are fundamental for the understanding of their mating patterns and evolution. Two closely related South American Nothofagus species were compared with regard to the differentiation times and positions of staminate and pistillate flowers along their parent growth units (GUs) by quantitative means.

Methods

Two samples of GUs that had extended in the 2004–2005 growing season were taken in 2005 and 2006 from trees in the Lanín National Park, Patagonia, Argentina. For the first sample, axillary buds of the parent GUs were dissected and the leaf, bud and flower primordia of these buds were identified. The second sample included all branches derived from the parent GUs in the 2005–2006 growing season.

Key Results

Both species developed flowering GUs with staminate and/or pistillate flowers; GUs with both flower types were the most common. The position of staminate flowers along GUs was similar between species and close to the proximal end of the GUs. Pistillate flowers were developed more distally along the GUs in N. alpina than in N. obliqua. In N. alpina, the nodes bearing staminate and pistillate flowers were separated by one to several nodes with axillary buds, something not observed in N. obliqua. Markovian models supported this between-species difference. Flowering GUs, including all of their leaves and flowers were entirely preformed in the winter buds.

Conclusions

Staminate and pistillate flowers of N. alpina and N. obliqua are differentiated at precise locations on GUs in the growing season preceding that of their antheses. The differences between N. alpina and N. obliqua (and other South American Nothofagus species) regarding flower distribution might relate to the time of anthesis of each flower type and, in turn, to the probabilities of self-pollination at the GU level.Key words: Branch, bud, growth unit, Markovian models, Nothofagus alpina, N. obliqua, Patagonian forests, pistillate flower, preformation, staminate flower     

Ectomycorrhizal fungal communities in Nothofagus nervosa (Raulí): A comparison between domesticated and naturally established specimens in a native forest of Patagonia,Argentina

Due to its overexploitation during the past century, Nothofagus nervosa is currently included in conservation and domestication programs, in which ectomycorrhizas play an important role. We aimed to describe the abundance and diversity of ectomycorrhizal fungi (EcMF) in both domesticated and naturally established N. nervosa specimens, and to analyse the influence of age, seasonality and forest management on EcMF communities. The occurrence of arbuscular mycorrhizas (AM) and dark septate endophytes (DSE) was also investigated. Fungal diversity and taxonomic identification were assessed by morphotyping and subsequent ITS-rDNA sequencing. Plant age, seasonality and forest management influenced EcMF communities. Colonization rates were higher than 90 % in all the specimens, and were significantly higher in mature trees and in autumn. The highest EcMF richness and diversity values were registered in domesticated specimens and in autumn. Most EcMF were basidiomycetes, belonging mainly to the Cortinariaceae and Tricholomataceae. Arbuscular mycorrhizas were not detected, while DSE were present within N. nervosa roots. Our results and previously published reports showed that some EcMF are capable of colonizing different Nothofagus species. In addition, the EcMF described in natural ecosystems are different from those colonizing N. nervosa during its cultivation in the nursery. These results improve our understanding of key factors affecting EcMF communities associated with Nothofagus in native forests and nurseries (age, season, forest management, cultivation techniques), and this information is relevant for improving domestication programs.     

Litter retention in Tasmanian headwater streams after clear-fell logging

Clear-fell logging around small headwater streams in Tasmanian wet eucalypt forests was predicted to affect both the retention of leaf litter and the composition and size of leaf packs. Retention structures were surveyed in six natural streams and six streams in forest regenerated 3–5 years after clear-fell and burn logging. Logged streams had more wood, but retained less leaves than natural streams, and consequently had fewer and smaller leaf packs. Leaf packs from natural streams contained 200% more leaves, bark and twigs than packs from logged streams. The effect of buoyancy on leaf retention was assessed with release and recapture of marked Eucalyptus obliqua and Nothofagus cunninghamii leaves. Eucalypt leaves were more likely to be trapped by retention structures on the bed of the stream, while smaller, more buoyant N. cunninghamii leaves were mainly trapped by leaf packs. Leaf packs in natural streams were formed on a matrix of small twigs and long strips of bark, shed from the upper branches of mature stringybark eucalypts, while leaf retention was reduced in logged streams because there are no mature trees to provide effective retention structures. Changes to the channel form increase both discharge and sedimentation. These factors have strong implications for downstream nutrient processing and riverine food webs.     

Comparative morphology of annual shoots in seedlings of five Nothofagus species from Argentinian Patagonia

Morphological features of seedlings were compared between five Nothofagus species from Patagonia: N. antarctica (Forst. F.) Oerst., N. dombeyi (Mirb.) Oerst, N. nervosa (Phil.) Dim. et Mil., N. obliqua (Mirb.) Oerst. and N. pumilio (Poepp. et Endl.) Krasser. Annual shoots of 1- to 3-year-old seedlings collected from natural communities were observed and the distribution of their length and number of internodes analysed. Each seedling produced one shoot each year. Leaves are alternate in all cases except for the cotyledons (which are always opposite) and the first two leaves after the cotyledons (which have, depending on the species, a high or low probability of being opposite to each other). Shoot length and number of internodes depended on the year and on the species concerned. Shoot length was more variable than internode number both within and between species. The probability of shoot apex death after shoot elongation may be high or low depending on the species and the age of the seedling.     

Mass loss and nutrient release from decomposing evergreen and deciduous Nothofagus litters from the Chilean Andes

Abstract Leaf litter decomposition experiments were conducted on two deciduous (Nothofagus obliqua (roble)) and Nothofagus pumilio (lenga)) and one evergreen (Nothofagus dombeyi) Nothofagus (Nothofagaceae) species from a single Chilean forest in order to understand how congeneric trees with differing leaf lifespans impact the soil in which they grow. Single‐species litter samples were decomposed in a mixed hardwood forest in Ohio and in a deciduous‐evergreen Nothofagus forest in Chile. In the Ohio forest, the two deciduous species’ litters decomposed at k ≈ 1.00 per year and the evergreen at k ≈ 0.75 per year. In Chile k ranged from k ≈ 0.06 (N. obliqua) to k ≈ 0.23 (N. pumilio) per year. In both experiments, N and P were released faster from the deciduous litters than from evergreen litter. In Ohio, evergreen litter immobilized more N and P for a longer time period than did deciduous litter. As N. dombeyi stands tend to have lower available soil N and P in this particular mixed Nothofagus forest, the increased time of N and P immobilization by N. dombeyi litter suggests a feedback role of the tree itself in perpetuating low N and P soil conditions.     

Pollination and self-interference in Nothofagus

Interference between male and female functions within a monoecious plant may hinder crossing and decrease seed set. We assessed the probability of self-pollination and the effect of self-pollination on cross-pollination for two self-incompatible species: Nothofagus obliqua and N. nervosa. The probability of self-pollination was studied by tracking the phenologies of staminate and pistillate flowers, including an analysis of stigmatic receptivity. Pure and mixed pollinations were performed in order to evaluate the effect of self-pollination upon cross-pollination. Phenological observations suggest that self-pollination is highly likely in both species. Compared to pure cross-pollination, the application of self-pollination prior to cross-pollination resulted in lower numbers of germinated pollen grains for both species, and also in a lower production of viable seeds in N. nervosa. The low proportion of viable seeds often observed in natural populations of N. obliqua and N. nervosa may be related to self-pollination.     

Bud and growth-unit structure in seedlings and saplings of Nothofagus alpina (Nothofagaceae)

In temperate trees, axis length growth generally results from the differentiation of organs at the end of a growing season and the extension of such "preformed organs" in the next growing season. Neoformation, i.e., the simultaneous differentiation and extension of organs, has been studied for only a few species. Here we evaluated bud composition and growth unit (GU) size for seedlings and saplings of Nothofagus alpina, a valuable South American forest tree. Trunk GUs of seedlings and saplings included preformed and neoformed organs, whereas main-branch GUs of saplings were entirely preformed. The size of a GU was more closely related to the number of preformed green leaves than to the number of cataphylls of its preceding bud. Proximal buds of a trunk GU had more cataphylls and less green-leaf primordia than distal buds. Individual leaf area increased from proximal to distal positions on trunk GUs. For trunk and main-branch GUs, the length/width ratio was maximum for leaves in intermediate positions. The development of large neoformed leaves at the end of the growing season could increase the photosynthetic capacity of this species in late summer, when the activity of preformed organs is likely to be decreasing.     

Effects of climate on the radial growth of mixed stands of Nothofagus nervosa and Nothofagus obliqua along a precipitation gradient in Patagonia,Argentina

Climate models for North Patagonia in Argentina project dryer conditions, due to a decrease in mean precipitation combined with an increase in mean temperatures. The temperate mixed Nothofagus forest, which exists along a steep precipitation gradient, could be directly impacted. For this study, we evaluated the influence of mean climate on the growth of the significant deciduous species: N. nervosa and N. obliqua. For the first time in Argentina, dendroclimatological analyses were done on both species using a network of 14 chronologies, covering their longitudinal distribution along a gradient of declining precipitation from west to east. Seasonal correlation analysis revealed that temperature has a negative effect on the growth of both species across all sites, particularly during summer of the previous and current growth season. Precipitation has a positive effect on the growth of trees for both species, which is more significant for N. nervosa. The relationship between early-summer climate and growth remained relatively stable over time for N. nervosa; however, for N. obliqua the detrimental effects of temperature increased towards the end of the 20th century, and the positive effects of precipitation decreased, particularly at the driest end of the gradient. These results suggest that a continued decrease in rainfall with a rise in temperature could impact growth for both of these species.     

Heteroblastic development and the optimal partitioning of traits among contrasting environments in Acacia implexa

Background and Aims: Optimal partitioning theory (OPT) predicts plants will allocatebiomass to organs where resources are limiting. Studies of OPTfocus on root, stem and leaf mass ratios where roots and stemsare often further sub-divided into organs such as fine roots/taproots or branches/main stem. Leaves, however, are rarely sub-dividedinto different organs. Heteroblastic species develop juvenileand adult foliage and provide an opportunity of sub-dividingleaf mass ratio into distinct organs. Acacia implexa (Mimosaceae)is a heteroblastic species that develops compound (juvenile),transitional and phyllode (adult) leaves that differ dramaticallyin form and function. The aims of the present study were togrow A. implexa to examine patterns of plastic development ofwhole-plant and leaf traits under the OPT framework. Methods: Plants were grown in a glasshouse under contrasting nutrient,light and water environments in a full factorial design. Allocationto whole-plant and leaf-level traits was measured and analysedwith multivariate statistics. Key Results: Whole-plant traits strongly followed patterns predicted by OPT.Leaf-level traits showed a more complex pattern in responseto experimental treatments. Compound leaves on low nutrientplants had significantly lower specific leaf area (SLA) andwere retained for longer as quantified by a significantly greatercompound leaf mass ratio after 120 d. There was no significantdifference in SLA of compound leaves in the light treatment,yet transitional SLA was significantly higher under the lowlight treatment. The timing of heteroblastic shift from compoundto transitional leaves was significantly delayed only in thelow light treatment. Therefore, plants in the light treatmentresponded at the whole-plant level by adjusting allocation toproductive compound leaves and at the leaf-level by adjustingSLA. There were no significant SLA differences in the watertreatment despite strong trends at the whole-plant level. Conclusion: Explicitly sub-dividing leaves into different types providedgreater insights into OPT.     

Nitrogen addition stimulates forest litter decomposition and disrupts species interactions in Patagonia,Argentina

Nitrogen (N) deposition and biodiversity loss are important drivers of global change, with uncertain consequences for carbon (C) and nutrient turnover in terrestrial ecosystems. We evaluated the simultaneous effects of N deposition and plant diversity on litter decomposition within a temperate forest in Patagonia. We identified ‘tree triangles’ created by the intersection of three tree‐canopies that directly controlled micro‐environmental conditions on the forest floor, and combined it with an N addition treatment. Triangles were composed of one or three Nothofagus species (N. dombeyi, N. obliqua or N. nervosa). We placed litterbags containing litter of each of the Nothofagus species and litterbags containing a mixture of the three species within all triangles and assessed mass loss over 2 years. We used a standard litter type in all triangles to independently evaluate triangle effects on decomposition. N addition had strong and positive effects on decomposition with an average 46% increase in the decomposition constant. Litter species significantly differed in their response to N addition; litter with higher lignin content and lower labile C content had larger increase in decomposition due to fertilization. Also, N addition disrupted two types of species interactions that control decomposition. The affinity relation between litter and decomposers, that enhanced decomposition of home litter (‘home‐field advantage’) that was demonstrated to be significant for all three Nothofagus species, disappeared with N addition. Second, N addition modified litter species interactions, transforming neutral effects of litter mixtures to positive, nonadditive effects on mass loss. Finally, N addition stimulated N release from decomposing litter which was modulated by plant species effects. Together, these results suggest that N addition to unpolluted forests increases C loss, contrary to what has been observed for temperate forests in industrialized areas of the world, and that alterations in nutrient pools have effects on species interactions, including the elimination of affinity effects for decomposition.     

Leaf functional traits of Neotropical savanna trees in relation to seasonal water deficit

The seasonal savannas (cerrados) of Central Brazil are characterized by a large diversity of evergreen and deciduous trees, which do not show a clear differentiation in terms of active rooting depth. Irrespective of the depth of the root system, expansion of new foliage in deciduous species occurs at the end of the dry season. In this study, we examined a suite of leaf traits related to C assimilation, water and nutrients (N, P) in five deciduous and six evergreen trees that were among the dominant families of cerrado vegetation. Maximum CO₂ assimilation on a mass basis (A_mass) was significantly correlated with leaf N and P, and specific leaf area (SLA; leaf area per unit of leaf mass). The highest leaf concentrations of both nutrients were measured in the newly mature leaves of deciduous species at the end of the dry period. The differences in terms of leaf N and P between evergreen and deciduous species decreased during the wet season. Deciduous species also invested less in the production of non-photosynthetic leaf tissues and produced leaves with higher SLA and maintained higher water use efficiency. Thus, deciduous species compensated for their shorter leaf payback period by maintaining higher potential payback capacity (higher values of A_mass) and lower leaf construction costs (higher SLA). Their short leafless period and the capacity to flush by the end of the dry season may also contribute to offset the longer payback period of evergreen species, although it may involve the higher cost of maintaining a deep-root system or a tight control of plant water balance in the shallow-rooted ones.     

Spatial variation in sapwood area to leaf area ratio and specific leaf area within a crown of silver birch

Spatial variation in sapwood area to leaf area ratio (Huber value, HV) and specific leaf area (SLA) was examined in branches of closed-canopy trees of silver birch (Betula pendula Roth). HV increased basipetally within a crown and decreased with increasing branch order, but exhibited no significant radial trend along a primary branch. HV was primarily determined by branch position in a crown and branch diameter at the sampling point, being independent of the size of the tree and branch. Greater HV in the lower-crown branches is considered a means to mitigate differences in hydraulic transport capacity between the branches located in different canopy layers. Beside branch position and sampling location on a branch, SLA depended significantly on several other variables characterising tree and branch size. SLA increased basipetally within a crown and along a primary branch, but exhibited no significant trend with branch orders. Because height caused leaf area (A_L) to diminish more rapidly than leaf dry weight, A_L primarily determined the vertical variation in SLA.     

Similarities and gradients in growth unit branching patterns during ontogeny in "Fuji" apple trees: a stochastic approach

This study aims to explore and model the changes in growth unit (GU) branching patterns during tree ontogeny. The question was addressed in apple trees cv. "Fuji", by analysing the relative impact of GU length and within-tree position. The development of two 6-year-old trees was recorded over 6 years. The fate of axillary buds along each GU was represented as a sequence of symbols corresponding to five types of lateral growth: latent buds, short, medium, long, and floral lateral GUs. Based on an exploratory analysis of data and a priori hypotheses, a hidden semi-Markov chain was estimated from all of these GU sequences. This model was composed of six transient states representing successive branching zones along the GUs. The accuracy of this global model was a posteriori assessed by fitting the characteristic distributions computed from model parameters to the corresponding empirical characteristic distributions extracted from the observed sequences. The observed sequences were then grouped hierarchically according to the GU length, year of growth, and branching order. Comparing model parameters between these sub-groups revealed similarities between GUs. These similarities were based on particular branching zones whose composition and relative position within the GUs remained invariant across the subgroups: the latent zones, floral zone, and short-lateral zone. The probability of occurrence of the floral zone varied with the year, showing the alternate fruiting of "Fuji". It is shown that, during tree ontogeny, as GU length decreases, branching patterns tend to progressively simplify due to the disappearance of the most central zones and a progressive reduction in the length of the floral zone.     

Ectomycorrhizal fungal communities associated to Nothofagus species in Northern Patagonia

Ectomycorrhizal fungi constitute an important component of soil biota in Nothofagus forests in Patagonia. However, ectomycorrhizal fungal community is poorly known in this region. Here, we assess biodiversity and community compositions of ectomycorrhizal fungal species associated with Nothofagus dombeyi, N. obliqua and N. alpina. We selected three monospecific Nothofagus forest sites for each species within the boundaries of the Lanin National Park in Northern Patagonia. Ectomycorrhizal fungal species were identified based on morphotyping and rDNA (ITS and 28S rDNA) sequence analysis using both universal and taxon-specific primers. Contrary to previous studies on congeneric host trees, our results showed no significant differences among Nothofagus forest types in terms of fungal biodiversity and community composition. However, altitude had a strong effect on the structure of the ectomycorrhizal fungal community associated with Nothofagus spp.     

Physiological and morphological short-term responses to light and temperature in two Nothofagus species of Patagonia, South America

The study of plant responses to environmental stress factors is essential for management of plant systems and for anticipating their response to climate change. The main goal of this study was to determine morphological and physiological responses of Nothofagus obliqua and N. nervosa seedlings to light and temperature, two of the main stress factors acting in their current natural distribution in NW Patagonia. Responses to light were evaluated analyzing growth and survival, as well as morphological and physiological traits related to them, in seedlings subjected to three contrasting light conditions (full-sun conditions, 50% of sunlight and 20% of sunlight) during one growth season. Temperature photosynthetic responses were evaluated in seedlings subjected to temperature treatments between ?5 and 40°C for 2 and 4 h. Growth rate and biomass partition were similar between light treatments in both species. High apical meristem damage and decreased photosynthetic capacity of preformed leaves were observed under full-sun conditions, suggesting that high light levels have a deleterious effect on plant yield. Both species produced neoformed leaves during the growing season with better photosynthetic capacity than preformed leaves under full sun conditions, contributing to plant acclimation. Almost no plasticity was observed in morphological traits in response to shade. Both species differed in optimum temperature for photosynthesis, with a wider temperature range at which high photosynthesis is maintained in N. obliqua. In both species the higher values of net photosynthetic rate were found at higher temperatures than the mean annual temperature of its current natural distribution range. Under no water-stress conditions, future higher temperatures could increase carbon fixation of these species, with a little advantage of N. obliqua if temperature variance is high. Synergy effect of various environmental stress factors, particularly considering cultivation of these species outside their current natural distribution sites require further studies.     

Integrating genetics and suitability modelling to bolster climate change adaptation planning in Patagonian <Emphasis Type="Italic">Nothofagus</Emphasis> forests

We investigated the impact of past changes in habitat suitability on the current patterns of genetic diversity of two southern beeches (Nothofagus nervosa and Nothofagus obliqua) in their eastern fragmented range in Patagonian Argentina, and model likely future threats to their population genetic structure. Our goal was to develop a spatially-explicit strategy for guiding conservation and management interventions in light of climate change. We combined suitability modelling under current, past (Last Glacial Maximum ~ 21,000 bp), and future (2050s) climatic conditions with genetic characterization data based on chloroplast DNA, isozymes, and microsatellites. We show the complementary usefulness of the distribution of chloroplast haplotypes and locally common allelic richness calculated from microsatellite data for identifying the locations of putative glacial refugia. Our findings suggest that contemporary hotspots of genetic diversity correspond to convergence zones of different expansion routes, most likely as a consequence of admixture processes. Future suitability predictions suggest that climate change might differentially affect both species. All genetically most diverse populations of N. nervosa and several of N. obliqua are located in areas that may be most severely impacted by climate change, calling for forward-looking conservation interventions. We propose a practical spatially- explicit strategy to target conservation interventions distinguishing priority populations for (1) in situ conservation (hotspots of genetic diversity likely to remain suitable under climate change), (2) ex situ conservation in areas where high genetic diversity overlaps with high likelihood of drastic climate change, (3) vulnerable populations (areas expected to be negatively affected by climate change), and (4) potential expansion areas under climate change.     

Herbivory alters plant carbon assimilation,patterns of biomass allocation and nitrogen use efficiency

Herbivory can trigger physiological processes resulting in leaf and whole plant functional changes. The effects of chronic infestation by an insect on leaf traits related to carbon and nitrogen economy in three Prunus avium cultivars were assessed. Leaves from non-infested trees (control) and damaged leaves from infested trees were selected. The insect larvae produce skeletonization of the leaves leaving relatively intact the vein network of the eaten leaves and the abaxial epidermal tissue. At the leaf level, nitrogen content per mass (N_mass) and per area (N_area), net photosynthesis per mass (A_mass) and per area (A_area), photosynthetic nitrogen-use efficiency (PNUE), leaf mass per area (LMA) and total leaf phenols content were measured in the three cultivars. All cultivars responded to herbivory in a similar fashion. The N_mass, A_mass, and PNUE decreased, while LMA and total content of phenols increased in partially damaged leaves. Increases in herbivore pressure resulted in lower leaf size and total leaf area per plant across cultivars. Despite this, stem cumulative growth tended to increase in infected plants suggesting a change in the patterns of biomass allocation and in resources sequestration elicited by herbivory. A larger N investment in defenses instead of photosynthetic structures may explain the lower PNUE and A_mass observed in damaged leaves. Some physiological changes due to herbivory partially compensate for the cost of leaf removal buffering the carbon economy at the whole plant level.     

Seedling drought stress susceptibility in two deciduous Nothofagus species of NW Patagonia

The physiological capacities of seedlings to cope with drought may be subject to strong selective pressure in the context of future climate scenarios, threatening the regeneration and sustainability of forests. Characterization of the responses and the variability between species is of interest to breeding and domestication programs. In this study, our main goal was to describe some of the physiological mechanisms involved in the drought response of Nothofagus nervosa and N. obliqua, two forest species of ecological and commercial importance (high wood quality) in NW Patagonia. We tested for differences in water status, gas exchange and survival in response to a gradually imposed severe drought. Based on cavitation vulnerability curves and hydraulic conductivity measurements, we can conclude that N. obliqua stems have higher specific hydraulic conductivity and somewhat lower vulnerability to cavitation than N. nervosa stems, leading it to sustain higher stomatal conductance under non-severe drought conditions. N. obliqua had higher photosynthetic capacity than N. nervosa, due both to characteristics of its hydraulic architecture and to its higher metabolic capacity. Our results indicate that both species present characteristics of plants susceptible to water stress. Also, both species showed behavior resembling an anisohydric response. This behavior results from a lack of stomatal control over transpiration while the soil dehydrates, probably accompanied by very high vulnerability to cavitation. In contrast, both species had similar high stomatal sensitivity to vapor pressure deficit when soil water was limiting.     

The role of leaf cellulose content in determining host plant preferences of three defoliating insects present in the Andean‐Patagonian forest

Phytophagous insects choose their feeding resources according to their own requirements in addition to properties of the host plants, such as biomechanical defences. The feeding preferences of the native folivorous insects of the Andean‐Patagonian forest (Argentina) have rarely been studied. These environments present a wide diversity and abundance of insects associated with trees of the Nothofagus and Lophozonia (Nothofagaceae) genera, which represent the main tree species of the forests of the southern hemisphere. In particular, Lophozonia alpina and Lophozonia obliqua are of great interest because they have a wide distribution, a high capacity for hybridization and exhibit great phenotypic plasticity. This versatility causes substantial variation in the biomechanical properties of leaves, affecting the feeding preferences of insects. The purpose of this work was to study the food selection behaviour of three leaf‐chewing insects (Polydrusus nothofagii, Polydrusus roseaus (Coleoptera: Curculionidae) and Perzelia arda (Lepidoptera: Oecophoridae)) associated with L. alpina and L. obliqua as host plants. Based on their choices, our aim was to determine a preference scale for each insect species and the variables on which these preferences were based. Therefore, we selected trees of L. alpina and L. obliqua, measured several properties such as cellulose content and recorded which leaves were eaten. As a result, we determined that the three species of insects feed on both host plants but prefer the leaves of L. obliqua, with cellulose content being the main determining factor for their decisions. However, in the case of P. arda, there was a positive relationship between cellulose and host plant preference, whereas there was an opposite relationship for the weevils. We conclude that during feeding selection, there are some properties of the leaves that have a more important role than others and that the same property does not exert the same behavioural response in all folivorous insects.