2. Plant growth and environment
2. Plant growth and environment
Light, CO2, temperature, water and nutrients are taken as key driving variables for growth
responses in a wide range of species. Growth indices, especially whole RGR, serve as an indicator of
plant response and of interactions between environmental factors where they occur. Variation in whole-
plant RGR is then resolved into contributions from NAR and LAR. Ecological implications for managed
and natural communities are considered.
Light
Light has an impact on both extent
and activity of plant canopies. Taking
cucumber as an archetype for herbaceous
crop plants (Figure 2) leaf growth increases
with daily irradiance due to increased cell
number rather than increased cell size. Leaf
thickness is also positively affected by daily
irradiance, principally resulting in a greater
depth of palisade (Table 1). Indeed, mean
cell volume is more than doubled under
strong irradiance (3.11 × 10–5 mm3 cf. 1.46
× 10–5 mm3 at 0.5 MJ m–2 d–1 because
cross-sectional area is virtually unchanged
cell depth is responsible. This greater depth
of palisade in strong light confers a greater
photo-synthetic capacity on such leaves
(expressed on an area basis) and translates
into larger values for NAR and a potentially
higher RGR. At lower irradiance (Table 1)
leaves are thinner and SLA will thus
increase with shading, and because LAR =
SLA × LWR a smaller absolute size at lower
irradiance can be offset by larger SLA
resulting in LAR increase.
Temperature
Temperature effects on RGR are generally
attributable to rate of canopy expansion rather
than rate of carbon assimilation. In the early days
of growth analysis, Blackman et al. (1955)
inferred from a multi-factor analysis of growth
response to environmental conditions that NAR
was relatively insensitive to temperature, but
whole-plant growth was obviously affected.
Using day/night temperature as a driving
variable, Potter and Jones (1977) provided a
detailed analysis of response in key growth
indices for a number of species (Table 2). Data for
maize, cotton, soybean, cocklebur, Johnson grass
and pigweed confirmed that 32/21°C was
optimum for whole-plant relative growth rate
(RGRW) as well as relative rate of canopy area
increase (RGRA). Both indices were lowest at
21/10°C. Moreover, variation in RGRW and
RGRA was closely correlated across species and
treatments (pooled data).
Carbon dioxide
Growth responses to elevated CO2 can
be spectacular, especially during early
exponential growth and derive largely
from direct effects of increased CO2
pressure on photosynthesis. C3 plants will
be most affected, and especially at high
temperature where photorespiratory loss
of carbon has the greatest impact on
biomass accumulation.
Nutrients (nitrogen and phosphorus)
Leaf expansion is particularly sensitive to nutrient supply (especially nitrogen, phosphorus,
potassium (N, P, K) and magnesium) due primarily to the needs of enlarging cells for synthesis of new
materials and generation of turgor. Nutrient deficiency or imbalance is first detected in leaf growth
rather than leaf assimilation, and in terms of canopy development, nutrient supply impacts on
phyllochron, RGR and fine leaf size. Plants grown on high, medium or low N supply (Table [N] 2.94%,
1.18% and 0.63% N (dry mass) respectively show a strong decline in final leaf size and RGR. While
phyllochron increased from high to low nutrient supply respectively.
Water
Growth is a turgor-dependent process enlargement are especially sensitive to water stress. When
plants encounter water stress, leaf area increase is either diminished or even ceases well ahead of any
clear reduction in leaf gas exchange. NAR is thus less sensitive to water stress than RGR.
Total plant biomass at final harvest was greatly reduced by the low in leaf expansion. NAR (area basis)
was not affected to the same extent as final biomass but NAR (‘protein’ basis) was substantially reduced
because leaf ‘protein’ was increased by water stress.
Takami et al. (1981) grew sunflowers in a greenhouse under natural light in Canberra (March–
May 1980). Seedlings were initially well watered to ensure good establi (first 15 d). After thinning to
two plants per pot, irrigation was then withheld some pots, and unstressed maintained near field
capacity. Drought stress developed slowly (as intended) and drought-stressed plants recovered fully
within 4–6 d of irrigation. Just prior to rewatering, pre-dawn leaf turgor was actually higher in stressed
plants (0.63 MPa) compared with controls (0.39) not with standing a rather lower bulk leaf water
potential (Ψleaf = MPa in stressed and control respectively).
Leaf growth dynamics (Table 8) are based on comparisons between control and stress-recovered
plants, and apply to corresponding nodes, namely 5, 13, 17 and 23. Final leaf size varies with node
number in sunflower, hence the need for strict correspondence. Leaves at node 5 (Table 8) encountered
an intensifying stress soon after appearance. Stressed plants maintained similar r, and failed to reach the
same final size (Ax) as well-watered controls. Taking r as indicative of cell division during the
exponential phase of lamina expansion with subsequent growth driven mainly by enlargement, drought
stress has restricted cell enlargement rather than cell division.
Leaves at node 13 on droughted plants (prior to stress relief on day 36) were similar in RGR (r =
0.24 d–1 cf. 0.26 d–1 in well-watered controls) but greatly restricted in final size (84 cm2 cf. 392 cm2 in
controls), again emphasising the sensitivity of cell enlargement to moisture stress.
Phyllochron (3t0 in Equation 6.14) was little affected up to node 13 (Table 8) but after-effects of
previous stress became apparent on rate of leaf appearance from node 14 to node 17, resulting in 3t0
increasing from 1.4 to 3.2 d. Subsequent leaf appearance (node 17 to node 23) was even accelerated in
stress-recovered plants, resulting in a Δt0 = 0.8 d cf. 1.1 d in non-stressed controls. r at node 23 was
unchanged by stress-recovery treatments but final size was substantially greater (228 cm2) in stress-
recovered compared with non-stressed controls (198 cm2). Such compensatory growth by individual
leaves following stress relief would draw on N-based resources that accumulate during drought, while
turgor-driven expansion to a greater final size could partly arise from drought-induced osmotic
adjustment.