Light impacts both photosynthetic activity and morphology of individual leaves and of plant canopies. Leaves are larger at higher light. Leaf area increases because of more cells per leaf rather than by cells having a larger surface area. Cells volume, however, increases and gives rise to substantial increases in leaf thickness. This is usually achieved by a greater depth of palisade cells, either greater in depth or an extra layer of cells.
In the example shown for cucumber in Table 6.1, high light caused a three-fold increase in area, but the cell cross-section was the same, indicating that the leaves had three times the number of cells. The cell volume more than doubled under high light (3.11 × 10–5 mm3 at 3.2 MJ m–2 s–1 cf. 1.46 × 10–5 mm3 at 0.5 MJ m–2 d–1), and because cross-sectional area was virtually unchanged, cell depth was responsible. This greater depth of palisade cells in strong light confers a greater photosynthetic capacity (per unit leaf area) and translates into larger values for NAR and a potentially higher RGR. At lower irradiance (Table 6.1) leaves are thinner and SLA will thus increase with shading. Because LAR = SLA × LWR (Equation 6.9) a smaller leaf area at lower irradiance is offset to some extent by a higher SLA for maximum light capture with the most efficient use of resources.
The signiﬁcance of LAR × NAR interaction for whole-plant growth was appreciated early by G.E. Blackman (Agriculture Dept, Oxford University), who in a series of papers analysed shade-driven growth responses for many species. RGR response to growing conditions in low light, and the degree to which upward adjustment in LAR could offset reduced NAR, was a recurring theme. In a series of 20 pot experiments, Blackman and Wilson (1951a) established a close relationship between NAR and daily irradiance where shade-dependent reduction in NAR was similar for 10 species. NAR was linearly related to log irradiance, and extrapolation to zero NAR corresponded to a light-compensation point of 6–9% full sun for eight species, and 14–18% full sun for two others. Signiﬁcantly, neither slope nor intercept differentiated sun-adapted plants such as barley, tomato, peas and sunflower from two shade-adapted species (Geum urbanum and Solanum dulcamara). LAR proved especially responsive to light and accounted for contrasts between sun plants and shade plants in their growth response to daily irradiance.
Concentrating on sunflower seedlings, Blackman and Wilson (1951b) conﬁrmed that NAR increased with daily irradiance (Figure 6.2) and that LAR was greatly decreased (Figure 6.2). Response in RGR reflected LAR and especially in young seedlings which also showed highest RGR and were most sensitive to shading. LAR appeared sensitive to both daily maxima as well as daily total irradiance.
A comparison between sunflower (Blackman and Wilson 1951b) and the woodland shade plant Impatiens parviflora (Evans and Hughes 1961) conﬁrms this principle of LAR responsiveness to irradiance (Figure 6.2). Sunflower achieved noticeably higher NAR in full sun than did I. parviflora, but LAR was considerably lower, and translated into a somewhat slower RGR for sunflower. This species contrast was much stronger in deep shade (12% full sun) where RGR for I. parviflora had fallen to 0.090 d–1 whereas sunflower was only 0.033 d–1. Clearly, I. parviflora is more shade tolerant, and retention of a faster RGR in deep shade is due both to greater plasticity in LAR as well as a more sustained NAR. Adjustments in both photosynthesis and respiration of leaves contribute to maintenance of higher NAR in shade-adapted plants growing at low irradiance.