Abstract

Within-canopy variation in light results in profound canopy profiles in foliage structural, chemical, and physiological traits. Studies on within-canopy variations in key foliage traits are often conducted in artificial environments, including growth chambers with only artificial light, and greenhouses with and without supplemental light. Canopy patterns in these systems are considered to be representative to outdoor conditions, but in experiments with artificial and supplemental lighting, the intensity of artificial light strongly deceases with the distance from the light source, and natural light intensity in greenhouses is less than outdoors due to limited transmittance of enclosure walls. The implications of such changes in radiation conditions on canopy patterns of foliage traits have not yet been analyzed. We developed model-based methods for retrospective estimation of distance vs. light intensity relationships, for separation of the share of artificial and natural light in experiments with combined light and for estimation of average enclosure transmittance, and estimated daily integrated light at the time of sampling (Qint,C), at foliage formation (Qint,G), and during foliage lifetime (Qint,av). The implications of artificial light environments were analyzed for altogether 25 studies providing information on within-canopy gradients of key foliage traits for 70 species × treatment combinations. Across the studies with artificial light, Qint,G for leaves formed at different heights in the canopy varied from 1.8- to 6.4-fold due to changing the distance between light source and growing plants. In experiments with combined lighting, the share of natural light at the top of the plants varied threefold, and the share of natural light strongly increased with increasing depth in the canopy. Foliage nitrogen content was most strongly associated with Qint,G, but photosynthetic capacity with Qint,C, emphasizing the importance of explicit description of light environment during foliage lifetime. The reported and estimated transmittances of enclosures varied between 0.27 and 0.85, and lack of consideration of the reduction of light compared with outdoor conditions resulted in major underestimation of foliage plasticity to light. The study emphasizes that plant trait vs. light relationships in artificial systems are not directly comparable to natural environments unless modifications in lighting conditions in artificial environments are taken into account.

Highlights

  • Light is a key environmental factor altering plant form and function (Givnish, 1988; Valladares, 2003; Niinemets, 2007; Pearcy, 2007; Poorter et al, 2009a)

  • There are extensive light gradients within-plant canopies and numerous studies have demonstrated that foliage structural, chemical, and physiological traits acclimate to these gradients (e.g., Hirose and Werger, 1987; Gutschick and Wiegel, 1988; Terashima and Hikosaka, 1995; Anten, 2005; Hikosaka, 2005; Niinemets, 2007), resulting in enhanced canopy carbon gain compared with invariable foliage characteristics (e.g., Niinemets and Anten, 2009; Dewar et al, 2012; Niinemets, 2012)

  • ACCLIMATION STUDIES IN ARTIFICIAL ENVIRONMENTS A number of influential pioneering studies on foliage acclimation to within-canopy light gradients have been conducted in artificial environments (e.g., Gutschick and Wiegel, 1988; Hirose et al, 1988; Schieving et al, 1992; Evans, 1993a,b; Pons et al, 1993; Pons and Jordi, 1998; Pons and Anten, 2004), and artificial environments remain an important platform to gain insight into plant acclimation (Table 1, Poorter et al, 2012)

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Summary

Introduction

Light is a key environmental factor altering plant form and function (Givnish, 1988; Valladares, 2003; Niinemets, 2007; Pearcy, 2007; Poorter et al, 2009a). There are extensive light gradients within-plant canopies and numerous studies have demonstrated that foliage structural, chemical, and physiological traits acclimate to these gradients (e.g., Hirose and Werger, 1987; Gutschick and Wiegel, 1988; Terashima and Hikosaka, 1995; Anten, 2005; Hikosaka, 2005; Niinemets, 2007), resulting in enhanced canopy carbon gain compared with invariable foliage characteristics (e.g., Niinemets and Anten, 2009; Dewar et al, 2012; Niinemets, 2012). Several classical studies investigating plant acclimation along light gradients have been conducted in artificial environments including greenhouses with natural illumination, greenhouses with natural, and supplemental illumination and growth chambers (e.g., Gutschick and Wiegel, 1988; Hirose et al, 1988; Evans, 1993a,b; Pons et al, 1993).

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