Seedling growth and biomass allocation in relation to leaf habit and shade tolerance among 10 temperate tree species

Initial growth of germinated seeds is an important life history stage, critical for establishment and succession in forests. Important questions remain regarding the differences among species in early growth potential arising from shade tolerance. In addition, the role of leaf habit in shaping relationships underlying shade tolerance-related differences in seedling growth remains unresolved. In this study we examined variation in morphological and physiological traits among seedlings of 10 forest tree species of the European temperate zone varying in shade tolerance and leaf habit (broadleaved winter-deciduous species vs needle-leaved conifers) during a 10-week period. Seeds were germinated and grown in a controlled environment simulating an intermediate forest understory light environment to resolve species differences in initial growth and biomass allocation. In the high-resource experimental conditions during the study, seedlings increased biomass allocation to roots at the cost of leaf biomass independent of shade tolerance and leaf habit. Strong correlations between relative growth rate (RGR), net assimilation rate (NAR), leaf area ratio (LAR), specific leaf area (SLA) and leaf mass fraction (LMF) indicate that physiology and biomass allocation were equally important determinants of RGR as plant structure and leaf morphology among these species. Our findings highlight the importance of seed mass- and seed size-related root morphology (specific root length - SRL) for shade tolerance during early ontogeny. Leaf and plant morphology (SLA, LAR) were more successful in explaining variation among species due to leaf habit than shade tolerance. In both broadleaves and conifers, shade-tolerant species had lower SRL and greater allocation of biomass to stems (stem mass fraction). Light-seeded shade-intolerant species with greater SRL had greater RGR in both leaf habit groups. However, the greatest plant mass was accumulated in the group of heavy-seeded shade-tolerant broadleaves. The results of our study suggest that the combinations of plant attributes enhancing growth under high light vary with shade tolerance, but differ between leaf habit groups

Plasticity in seedling morphology, biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle-leaved) and observed tolerance to shade, when growing in two contrasting light treatments - open (about 20% of full sunlight) and shade (about 5% of full sunlight). We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments. Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) - leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area-based rates of light-saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade. We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle-leaved conifers in response to shade. However, an expectation of higher plasticity in shade-intolerant species than in shade-tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit