Intraspecific Variation In Growth Research Articles

Using intraspecific molecular and phenotypic variation to promote multi-functionality of reforestation during climate change – A review of tropical forest case studies in South-east Asia

Abstract The study of intraspecific genetic variation in plant traits for use in tropical forest restoration has broad potential for increasing our ability to achieve multi-functional objectives during this era of climate change. Developing seed-sourcing guidelines that optimize phenotypic characteristics best suited to a particular planting site as well as to future conditions imposed by environmental change could be useful for effective reforestation. Because evolution operates differently across tree species, this is an especially cumbersome task in tropical forests that contain thousands of species. Partially due to this high plant diversity, research and application of intraspecific variation in genetics, plant traits, and plant function in tropical forests wane far behind less diverse forest biomes. To examine the potential for improving reforestation efforts in tropical forests by considering intraspecific variation in plant traits and functions, we review the state of knowledge on intraspecific variation in South-east Asia as a case study. We focus on the dipterocarp family ( Dipterocarpaceae ), a highly diverse family of 16 genera with approximately 695 known species that often dominate lowland tropical rainforests of South-east Asia with many of these forests in a degraded state and in need of restoration. We found that there is research accumulating to understand genetic variation in approximately 10% of these 695 species. Intraspecific molecular variation exists at different spatial scales among species with 74% of species having moderate to high population differentiation (Fst > 0.10) and 92% of species with evidence of fine-scale genetic structure. Although this suggests a high potential for trait variation, few studies associated molecular with phenotypic variation. Seventeen tree species across 11 studies revealed intraspecific variation in traits or functions. Research indicates that intraspecific variation in growth may vary two-fold and drought tolerance four-fold among genotypes highlighting the possibility to pre-adapt trees to climate change during reforestation and to use intraspecific variation to promote the use of native species in commercial forestry. Our review presents opportunities and ideas for developing seed-sourcing guidelines to take advantage of intraspecific variation in traits and function by identifying how to locate this variation, which species would benefit, and how to test for trait variation. We also highlight an emerging area of research on local adaptation, common garden studies, and adaptive drought conditioning to improve reforestation during climate change.

Infection Causes Trade-Offs between Development and Growth in Larval Amphibians.

AbstractTrade-offs between life history traits are context dependent; they vary depending on environment and life stage. Negative associations between development and growth often characterize larval life stages. Both growth and development consume large parts of the energy budget of young animals. The metabolic rate of animals should reflect differences in growth and developmental rates. Growth and development can also have negative associations with immune function because of their costs. We investigated how intraspecific variation in growth and development affected the metabolism of larval amphibians and whether intraspecific variation in growth, development, and metabolic rate could predict mortality and viral load in larvae infected with ranavirus. We also compared the relationship between growth and development before and after infection with ranavirus. We hypothesized that growth and development would affect metabolism and predicted that each would have a positive correlation with metabolic rate. We further hypothesized that allocation toward growth and development would increase ranavirus susceptibility and therefore predicted that larvae with faster growth, faster development, and higher metabolic rates would be more likely to die from ranavirus and have higher viral loads. Finally, we predicted that growth rate and developmental rate would have a negative association. Intraspecific variation in growth rate and developmental rate did not affect metabolism. Growth rate, developmental rate, and metabolism did not predict mortality from ranavirus or viral load. Larvae infected with ranavirus exhibited a trade-off between developmental rate and growth rate that was absent in uninfected larvae. Our results indicate a cost of ranavirus infection that is potentially due to both the infection-induced anorexia and the cost of infection altering priority rules for resource allocation.

Introduced populations of the garden lupine are adapted to local generalist snails but have lost alkaloid diversity

Intraspecific variation in growth and defence among plant populations can be driven by differences in (a)biotic conditions, such as herbivory and resources. Introduction of species to novel environments affects simultaneously herbivory encountered by a plant and resource availability both directly and via altered competitive environment. Here, we address the question of how growth (leaf mass per area (LMA), plant size) and resistance traits (leaf alkaloids, leaf trichomes, resistance to a generalist snail) vary and covary between native and introduced populations of the garden lupine, Lupinus polyphyllus. We focused specifically on evolved differences among populations by measuring traits from plants grown from seed in a common environment. Plants from the introduced populations were more resistant against the generalist snail, Arianta arbustorum, and they had more leaf trichomes and higher LMA than plants from the native populations. The composition of alkaloids differed between native and introduced populations, with the native populations having more diversity in alkaloids among them. Resistance was positively associated with plant size and LMA across all populations. Other trait associations differed between native and introduced areas, implying that certain trade-offs may be fundamentally different between native and introduced populations. Our results suggest that, for the introduced populations, the loss of native herbivores and the alterations in resource availability have led to a lower diversity in leaf alkaloids among populations and may facilitate the evolution of novel trait optima without compensatory trade-offs. Such phytochemical similarity among introduced populations provides novel insights into mechanisms promoting successful plant invasions.

Intraspecific Variation in Growth and Alkaline Phosphatase Activity of Cylindrospermopsis raciborskii Strains in Response to Different Phosphorus Concentrations and Sources

The cyanobacterial species C. raciborskii are ubiquitous in tropical regions, and its successful invasion into temperate zones has been partially attributed to its ability of survival in low P availability and the existence of multiple ecotypes. To explore the physiological response of different strains to phosphorus fluctuations, four strains of C. raciborskii isolated from the Zhenhai Reservoir were used to investigate their growth and alkaline phosphatase (ALP) activity at different inorganic phosphorus (Pi) concentrations (HP=7.13 mg ·L-1, MP=0.64 mg ·L-1, LP=0.03 mg ·L-1) and different phosphorus forms [dipotassium hydrogen phosphate (K2HPO4), sodium pyrophosphate (K4P2 O7), sodium polyphosphate (K5P3O10), D-glucose-6-phosphate (D-G-6-P), adenosine triphosphate (ATP), cyclic adenosine monophosphate (cAMP)]. Four C. raciborskii strains showed a similar growth response to phosphate changes: their biomass increased with an increase in Pi concentrations, while the ALP activity showed the opposite trend. The ALP activity of C. raciborskii N8 was significantly lower than that of other three strains, regardless of inorganic phosphorus concentrations, suggesting that this strain had a higher adaptability to phosphorus fluctuations. When cultured with different phosphorus forms, the biomass of C. raciborskii N8 and N9 in three dissolved inorganic phosphorus (DIP) compounds were significantly higher than those in three dissolved organic phosphorus (DOP) compounds, with the maximum and minimum specific growth rate in K2HPO4and ATP treatments, respectively. C. raciborskii preferred DIP although they can also utilize DOP to sustain its growth. Under the DOP conditions, the ALP activity of C. raciborskii N8 in the ATP treatment was significantly higher than that in the other two organic phosphorus compounds, while we did not observe similar results in C. raciborskii N9, indicating that strain N8 was more sensitive to DIP deficiency. Our results showed an intraspecific variation within C. raciborskii strains from the same reservoir. Compared with the other strains, strain N8 represented better adaptability to phosphorus fluctuations and DIP deficiency. Variations within C. raciborskii strains may make this species more adaptable to environmental changes and enhance its competitive advantage.

Intraspecific Variation in Growth and Reproduction of the Marine Leech, Heptacyclus buthi.

Heptacyclus buthi was harvested from fish hosts in rocky intertidal zones of Sonoma and Marin Counties, California, in October 2008 (n = 162) and October 2010 (n = 51). The size of the leeches was quantified using a method that approximated the sagittal cross-section of each specimen. Size-frequency curves were modeled to estimate the number of size-class cohorts in each year. If H. buthi is an annual species like many of its relatives, the single cohort modeled for in 2010 and the comparable "older" cohort in 2008, both with a broad range of sizes, may represent 1 component of its reproductive life history. A second, younger, more-numerous, less-variable cohort modeled from the 2008 sample may represent a second reproductive bout during that year that was prevented in the subsequent La Niña period of 2010-2011.

Intra-specific variation in growth and wood density traits under water-limited conditions: Long-term-, short-term-, and sudden responses of four conifer tree species

Consequences of climate change will severely affect forest ecosystems in the near future, yet our understanding of how and why trees are responding to their abiotic environment is still limited. Intra-specific variation (ITV) in the growth response of trees to warming and drought has been widely neglected so far, but could play a key role for adapting forests to future climate conditions. We analyzed tree rings from four conifers (Picea abies, Abies alba, Larix decidua, Pseudotsuga menziesii) regarding their intra-specific adaptation potential when trees are growing at the warm and dry margins of species distributions. Our study comprises data from four common garden experiments (45 provenances and a total of 743 trees) and assessed growth response at different temporal scales from decades (long-term) to only a few event years (short-term) and finally for density fluctuations within one year (sudden response). We observed significant variation among provenances at all time-scales, but with varying degree among species. However, variation in short-term response (drought years) was remarkably unstable across all species, when the seasonal variation of drought occurrence was considered. Silver-fir and Douglas-fir showed significant associations between seed-source climate and growth response as well as trade-offs between early- and latewood growth reaction which strongly suggests that growth responses are adaptive. Intra-specific variation in conifers in response to drought will probably be sufficient to mitigate climate change consequences on forest growth, but growth-environment interactions as well as dependencies between temporal scales could create major pitfalls for adaptive forest management in the future.

Involvement of insulin-like growth factor in intraspecific variation in growth of Pacific oyster Crassostrea gigas during winter

Insulin-like growth factor I (IGF-I) plays a critical role in regulating proliferation, differentiation, growth, and survival in fish as well as in humans. Few studies have investigated the involvement of IGFs in bivalves. In Korea, the Pacific oyster Crassostrea gigas is an important aquaculture species, and its growth is dependent on environmental conditions. Intrinsic factors that can be used as indicators to assess oyster growth based on individual size are limited. This study analyzed IGF-I, IGF-I receptor (IGF-IR), and IGF binding protein (IGFBP) in adductor muscle of C. gigas in winter using immunoblotting and reverse transcriptase-polymerase chain reaction. Levels of IGF-I and the IGF-IR β subunit, and expression of C. gigas insulin receptor-related receptor and IGFBP messenger RNA were significantly higher in large individuals (LI) than small individuals (SI). The IGF-IR β subunit was detected in the adductor muscle of both groups; however, phosphorylation of the IGF-IR β subunit and extracellular signal-regulated kinase was greater in LI than SI. The condition index of C. gigas showed a similar pattern. The results of this study suggest that the IGF system is associated with growth rate differences in C. gigas during winter in Korea.

When mutualisms matter: Rhizobia effects on plant communities depend on host plant population and soil nitrogen availability

Abstract Mutualistic interactions, such as the relationship between legumes and rhizobia, can affect community properties, yet there needs to be greater understanding of when these interactions may be most important to communities. Resource mutualism theory provides predictions based on variation in abiotic and biotic factors. First, there is substantial intraspecific genetic variation in how legumes interact with rhizobia, including variation in legume growth response to rhizobia, number of rhizobia‐housing nodules and nitrogen fixation. Rhizobia likely have greater community impacts when associating with hosts that benefit greatly from rhizobia, produce many nodules and fix a lot of nitrogen. Second, increased soil nutrient availability typically reduces the partner benefits of legume–rhizobium interactions. As a result, the legume–rhizobium resource mutualism may have greater effects on communities in low‐nitrogen environments. We first conducted a common garden study of the annual legume Chamaecrista fasciculata to explore intraspecific variation in focal traits related to competition and mutualistic interactions with rhizobia. Then, we manipulated the presence of rhizobia, nitrogen fertilization and population identity of C. fasciculata in mesocosms simulating native plant communities. We measured plant diversity, assessed changes in community composition and measured soil nutrient availability. We detected significant genetically based intraspecific variation in growth and mutualism‐related traits across C. fasciculata populations. From the experimental mesocosms, we found that although rhizobia consistently reduced diversity, rhizobia most strongly reduced diversity when associating with C. fasciculata populations that were highly dependent on rhizobia and when nitrogen was most limiting. We also detected greater variation among populations in their effects on communities in the absence of rhizobia; the presence of rhizobia dramatically minimized effects of population identity on the surrounding community. Synthesis. These findings show that abiotic environmental factors and intraspecific variation in a dominant host plant can influence the magnitude of mutualism effects on communities and, reciprocally, how the presence of key mutualists can influence the importance of genetically based variation among populations to community outcomes. While prior studies have detected both strong and weak effects of mutualism on plant communities, our study illustrates that mutualism theory may predict when strong mutualism effects are most likely to occur and also shows how theories developed for particular species interactions may make studies investigating the ecological impacts of genetic variation a more predictive science.

Potassium starvation affects biomass partitioning and sink–source responses in three sweet potato genotypes with contrasting potassium-use efficiency

Rooted single leaves of sweet potato (Ipomoea batatas L.) produce and translocate photosynthates, thus representing an ideal model for characterising the source–sink relationships and responses to various environments. A hydroponics culture study was conducted with rooted single leaves of sweet potato to determine intraspecific variation in growth, biomass partitioning, and associated physiological changes in response to variable potassium (K) supply among genotypes Ji22 (low K-use efficiency), Nan88 (high K-uptake efficiency) and Xu28 (high K-use efficiency). Potassium deficiency suppressed biomass accumulation in blades, petioles and roots in all three genotypes. Root length of diameters <0.25 mm and 0.25–0.5 mm was significantly less for K-deficient than K-sufficient roots of all genotypes, but the difference was proportionally greater in the K-inefficient genotype Ji22 than the other two genotypes. Potassium deficiency also severely inhibited net photosynthesis of blades in Nan88 and Ji22, as well reducing photosynthate translocation, increasing starch, hexose and sucrose concentrations, and decreasing K concentration in blades. The genotypes varied in photosynthesis-related responses to the K deficiency. Xu28 had greater blade K concentration and net photosynthesis as well as stable maximum quantum yield of photosystem II (FV/FM, with FV = FM – F0) under K deficiency, possibly because of a better source–sink balance and more efficient translocation of photosynthates to roots and K to blade compared with genotypes Ji22 and Nan88. Impaired phloem loading during K deficiency was associated with a decline in photosynthetic rate and decreased carbohydrate supply from blades, resulting in restricted root growth.

Species characteristics and intraspecific variation in growth and photosynthesis of Cryptomeria japonica under elevated O3 and CO2.

In order to predict the effects of future atmospheric conditions on forest productivity, it is necessary to clarify the physiological responses of major forest tree species to high concentrations of ozone (O3) and carbon dioxide (CO2). Furthermore, intraspecific variation of these responses should also be examined in order to predict productivity gains through tree improvements in the future. We investigated intraspecific variation in growth and photosynthesis of Cryptomeria japonica D. Don, a major silviculture species in Japan, in response to elevated concentrations of O3 (eO3) and CO2 (eCO2), separately and in combination. Cuttings of C. japonica were grown and exposed to two levels of O3 (ambient and twice-ambient levels) in combination with two levels of CO2 (ambient and 550 µmol mol-1 in the daytime) for two growing seasons in a free-air CO2 enrichment experiment. There was no obvious negative effect of eO3 on growth or photosynthetic traits of the C. japonica clones, but a positive effect was observed for annual height increments in the first growing season. Dry mass production and the photosynthetic rate increased under eCO2 conditions, while the maximum carboxylation rate decreased. Significant interaction effects of eO3 and eCO2 on growth and photosynthetic traits were not observed. Clonal effects on growth and photosynthetic traits were significant, but the interactions between clones and O3 and/or CO2 treatments were not. Spearman's rank correlation coefficients between growth traits under ambient conditions and for each treatment were significantly positive, implying that clonal ranking in growth abilities might not be affected by either eO3 or eCO2. The knowledge obtained from this study will be helpful for species selection in afforestation programs, to continue and to improve current programs involving this species, and to accurately predict the CO2 fixation capacity of Japanese forests.

The southern South American Proteaceae, Embothrium coccineum exhibits intraspecific variation in growth and cluster-root formation depending on climatic and edaphic origins

Background and aims Cluster root (CR) functioning has been studied mainly in Proteaceae species from (P)-deficient old soils. However, in southern South America, six species occur in young P rich soils. The aims were: i) to study the growth and CR formation of Embothrium coccineum seedlings from populations contrasting in edaphic and climatic conditions and, ii) to study the effect of P availability on CR formation.

Intraspecific variation in leaf growth of wheat (Triticum aestivum) under Australian Grain Free Air CO2 Enrichment (AGFACE): is it regulated through carbon and/or nitrogen supply?

Underlying physiological mechanisms of intraspecific variation in growth response to elevated CO2 concentration [CO2] were investigated using two spring wheat (Triticum aestivum L.) cultivars: Yitpi and H45. Leaf blade elongation rate (LER), leaf carbon (C), nitrogen (N) in the expanding leaf blade (ELB, sink) and photosynthesis (A) and C and N status in the last fully expanded leaf blade (LFELB, source) were measured. Plants were grown at ambient [CO2] (~384µmolmol-1) and elevated [CO2] (~550µmolmol-1) in the Australian Grains Free Air CO2 Enrichment facility. Elevated [CO2] increased leaf area and total dry mass production, respectively, by 42 and 53% for Yitpi compared with 2 and 13% for H45. Elevated [CO2] also stimulated the LER by 36% for Yitpi compared with 5% for H45. Yitpi showed a 99% increase in A at elevated [CO2] but no A stimulation was found for H45. There was a strong correlation (r2=0.807) between LER of the ELB and soluble carbohydrate concentration in LFELB. In ELB, the highest spatial N concentration was observed in the cell division zone, where N concentrations were 67.3 and 60.6mg g-1 for Yitpi compared with 51.1 and 39.2mg g-1 for H45 at ambient and elevated [CO2]. In contrast, C concentration increased only in the cell division and cell expansion zone of the ELB of Yitpi. These findings suggest that C supply from the source (LFELB) is cultivar dependent and well correlated with LER, leaf area expansion and whole-plant growth response to elevated [CO2].

Intraspecific variation in the growth and survival of juvenile fish exposed to Eucalyptus leachate

Whilst changes in freshwater assemblages along gradients of environmental stress have been relatively well studied, we know far less about intraspecific variation to these same stressors. A stressor common in fresh waters worldwide is leachates from terrestrial plants. Leachates alter the physiochemical environment of fresh waters by lowering pH and dissolved oxygen and also releasing toxic compounds such as polyphenols and tannins, all of which can be detrimental to aquatic organisms. We investigated how chronic exposure to Eucalyptus leaf leachate affected the growth and survival of juvenile southern pygmy perch (Nannoperca australis) collected from three populations with different litter inputs, hydrology and observed leachate concentrations. Chronic exposure to elevated leachate levels negatively impacted growth and survival, but the magnitude of these lethal and sublethal responses was conditional on body size and source population. Bigger fish had increased survival at high leachate levels but overall slower growth rates. Body size also varied among populations and fish from the population exposed to the lowest natural leachate concentrations had the highest average stress tolerance. Significant intraspecific variation in both growth and survival caused by Eucalyptus leachate exposure indicates that the magnitude (but not direction) of these stress responses varies across the landscape. This raises the potential for leachate-induced selection to operate at an among-population scale. The importance of body size demonstrates that the timing of leachate exposure during ontogeny is central in determining the magnitude of biological response, with early life stages being most vulnerable. Overall, we demonstrate that Eucalyptus leachates are prevalent and potent selective agents that can trigger important sublethal impacts, beyond those associated with more familiar fish kills, and reiterate that dissolved organic carbon is more than just an energy source in aquatic environments.

Genotypic variation in traits linked to climate and aboveground productivity in a widespread C4 grass: evidence for a functional trait syndrome

Examining intraspecific variation in growth and function in relation to climate may provide insight into physiological evolution and adaptation, and is important for predicting species responses to climate change. Under common garden conditions, we grew nine genotypes of the C₄ species Panicum virgatum originating from different temperature and precipitation environments. We hypothesized that genotype productivity, morphology and physiological traits would be correlated with climate of origin, and a suite of adaptive traits would show high broad-sense heritability (H(2)). Genotype productivity and flowering time increased and decreased, respectively, with home-climate temperature, and home-climate temperature was correlated with genotypic differences in a syndrome of morphological and physiological traits. Genotype leaf and tiller size, leaf lamina thickness, leaf mass per area (LMA) and C : N ratios increased with home-climate temperature, whereas leaf nitrogen per unit mass (Nm ) and chlorophyll (Chl) decreased with home-climate temperature. Trait variation was largely explained by genotypic differences (H(2) = 0.33-0.85). Our results provide new insight into the role of climate in driving functional trait coordination, local adaptation and genetic divergence within species. These results emphasize the importance of considering intraspecific variation in future climate change scenarios.

Intraspecific variation in growth and yield response to elevated CO2 in wheat depends on the differences of leaf mass per unit area.

In order to investigate the underlying physiological mechanism of intraspecific variation in plant growth and yield response to elevated CO2 concentration [CO2], seven cultivars of spring wheat (Triticum aestivum L.) were grown at either ambient [CO2] (~384μmolmol-1) or elevated [CO2] (700μmolmol-1) in temperature controlled glasshouses. Grain yield increased under elevated [CO2] by an average of 38% across all seven cultivars, and this was correlated with increases in both spike number (productive tillers) (r=0.868) and aboveground biomass (r=0.942). Across all the cultivars, flag leaf photosynthesis rate (A) increased by an average of 57% at elevated [CO2]. The response of A to elevated [CO2] ranged from 31% (in cv. H45) to 75% (in cv. Silverstar). Only H45 showed A acclimation to elevated [CO2], which was characterised by lower maximum Rubisco carboxylation efficiency, maximum electron transport rate and leaf N concentration. Leaf level traits responsible for plant growth, such as leaf mass per unit area (LMA), carbon (C), N content on an area basis ([N]LA) and the C:N increased at elevated [CO2]. LMA stimulation ranged from 0% to 85% and was clearly associated with increased [N]LA. Both of these traits were positively correlated with grain yield, suggesting that differences in LMA play an important role in determining the grain yield response to elevated [CO2]. Thus increased LMA can be used as a new trait to select cultivars for a future [CO2]-rich atmosphere.

Intraspecific variation in growth and allocation patterns in seedlings of Pinus pinaster Ait. submitted to contrasting watering regimes: can water availability explain regional variation?

• Interpopulation variation in key functional traits of Pinus pinaster Ait. is well recognized. However, the relative importance of drought tolerance to explain this regional variation in the species remains elusive. • Here, we raise the question whether water availability constitutes a likely driver of regional variation in biomass allocation, growth and morphological traits of ten populations that cover the distribution range of P. pinaster. We carried out an experiment where seedlings of five families per population were submitted to two contrasting watering treatments. • The effects of water availability and population were significant for relative diameter and height growth rate, biomass allocation and number of lateral stems and dwarf shoots. Total dry mass significantly differed between watering treatments but it did not among populations. Populations could be clustered into four main groups. Root mass fraction explained most of the variation and significantly correlated to altitude but not to aridity. • The geographical pattern of genetic variation found in morphology and biomass allocation did not translate into population differences in drought tolerance or phenotypic plasticity to water availability, indicating that water availability is not a likely driver of the regional variation observed in the studied traits of P. pinaster at the seedling stage. Mots-cles : altitude / tolerance a la secheresse / Bassin mediterraneen / modeles mixtes / performance du plant entier

Intraspecific Variation in Growth of Marsh Macrophytes in Response to Salinity and Soil Type: Implications for Wetland Restoration

Genetic diversity within plant populations can influence plant community structure along environmental gradients. In wetland habitats, salinity and soil type are factors that can vary along gradients and therefore affect plant growth. To test for intraspecific growth variation in response to these factors, a greenhouse study was conducted using common plants that occur in northern Gulf of Mexico brackish and salt marshes. Individual plants of Distichlis spicata, Phragmites australis, Schoenoplectus californicus, and Schoenoplectus robustus were collected from several locations along the coast in Louisiana, USA. Plant identity, based on collection location, was used as a measure of intraspecific variability. Prepared soil mixtures were organic, silt, or clay, and salinity treatments were 0 or 18 psu. Significant intraspecific variation in stem number, total stem height, or biomass was found in all species. Within species, response to soil type varied, but increased salinity significantly decreased growth in all individuals. Findings indicate that inclusion of multiple genets within species is an important consideration for marsh restoration projects that include vegetation plantings. This strategy will facilitate establishment of plant communities that have the flexibility to adapt to changing environmental conditions and, therefore, are capable of persisting over time.

Habitat related growth and reproductive investment in estuarine waters, illustrated for the tellinid bivalve Macoma balthica (L.) in the western Dutch Wadden Sea

In estuarine areas, bivalve species can be found in a variety of environments, where they experience large differences in environmental conditions. In the present paper, the importance of different habitats (intertidal, subtidal and adjacent coastal waters) for the persistence of the population was evaluated for the bivalve Macoma balthica (L.) in the western Dutch Wadden Sea estuary. Intra-specific variation in growth and reproductive output were followed during the year and related to local abiotic conditions. Significant differences in growth and reproductive investment were found between locations. Young individuals were mostly found in the intertidal area, where growth in terms of somatic mass was good. These areas were not favourable for adult individuals, since growth in shell length was low and many individuals did not reproduce. In the subtidal, where the highest densities were found, somatic and gonadal mass indices were low. Coastal areas had the lowest densities and showed high growth in terms of shell length and body mass. The habitat with the highest reproductive effort per individual was not the most important habitat in terms of reproductive output due to differences in density and in size of the habitat type. For M. balthica, the subtidal habitat contributed the most to the reproductive output of the western Dutch Wadden Sea population although the highest reproductive output per individual was in the coastal area.

Carbon availability affects nitrogen source utilisation by Hymenoscyphus ericae

We compared the ability of five strains of the ericoid mycorrhizal fungus Hymenoscyphus ericae to utilise glutamine, ammonium or nitrate at high or low carbon (C) availability. The pattern of intraspecific variation in growth was affected by C availability. When C supply was high, growth differences between strains were explained by the total amount of nitrogen (N) taken up, suggesting variation in uptake kinetics. Under C-limiting conditions, strain differences were linked with their nitrogen use efficiency, implying intraspecific differences in N metabolism. The relationship between growth on glutamine and pH shifts in the media indicated that there was intraspecific variation in glutamine transporters. In addition, the correlation between pH changes and the amount of glutamine-N recovered as ammonium in the media indicated that there were intraspecific variations within the enzymatic pathways involved in glutamine metabolism. Our findings, compared with those of a previous study involving the same ericoid strains, draw attention to the temporal variation in nitrogen source utilisation by ericoid mycorrhizal fungi when maintained in axenic culture.

Intraspecific variation in growth, defense and herbivory in Dialium guianense (Caesalpiniaceae) mediated by edaphic heterogeneity

Based on resource allocation theory, a negative correlation is predicted between resource availability and plant defense against herbivore attack. Plants growing in resource-limited environments should display lower growth and higher defense against herbivores than plants growing where resources are less limited. Interspecific comparisons generally support these predictions. We evaluated this hypothesis at the intraspecific level, for two sapling populations of the canopy tree Dialium guianense (Caesalpiniaceae) at the Lacandona rain forest in southeast Mexico. The two populations occur in nearby sites, adjacent to the Chajul Field Station, under the same climatic conditions and within the same vegetation type, but with considerable differences in soil quality. The Floodplain site, under the influence of the Lacantun River, has favorable conditions for plant growth, in terms of nutrient and water availability, whereas the Hills site, given its location and soil characteristics, provides more restricted conditions for plant growth. Plants in the Floodplain site had higher growth and lower concentration of phenolic compounds than plants in the Hills (a two-fold difference in leaf area production, 1.3 less total phenolics). These differences were correlated with differences in herbivore attack, as saplings from the Hills, with a higher defensive potential, had lower average levels of herbivory than Floodplain plants (3.86% ± 0.80 vs. 7.75% ± 1.43 of leaf area loss). The relationship between the concentration of phenolic compounds and leaf quality for herbivores was consistent with preference assays carried out under laboratory conditions using two species of generalist herbivores, the army worm Spodoptera fugiperda and the native katydid Orophus sp. In 63.8 and 81.3% of the cases, third-instar larvae of S. fugiperda and adults of Orophus, respectively, preferred leaflets from the Floodplain plants population. Moreover, on average, the adults of Orophus consumed 2.9 times more leaf area from the Floodplain than from the Hills. In addition, a reciprocal transplant experiment indicated that phenotypic plasticity is likely to be the mechanism by which the plants expressed differential growth and traits affecting herbivory levels. In this experiment, growth and herbivory levels were 1.6 and 1.7 times higher, respectively, in plants transplanted into a Floodplain experimental plot than those in a Hills plot. This work contributes to our understanding of how edaphic heterogeneity can determine intraspecific variation in the relationship of plants with their herbivores and evaluates the underlying mechanisms promoting such influence.