CHAPTER 3: ALLOMETRY AND BIOMASS RESULTS AND DISCUSSION
3.8 Intercepted radiation and growth
Photosynthetically active radiation consists of wavelengths of solar radiation that are utilized by plant biochemical processes in photosynthesis to convert light energy into biomass. The light energy in turn can be converted into the number of photons of light intercepted by foliage, which, in forests, is directly related to the quantity of carbon assimilated from the atmosphere (Landsberg and Waring, 1997). Approximately 47% of this assimilated carbon is lost through respiration and the remainder is partitioned between leaves, roots and woody biomass (Landsberg and Waring, 1997, Waring, 1998). Partitioning between stand components may be influenced (within limits) by growth resource availability. Itfollows that there should be some relationship between absorbed PAR and AGB produced (Linder, 1985; Beadle, 1997).
Forest growth may be linearly related to light interception, allowing for the conelation of growth with LAI or light interception (McMUltrie et aI., 1994; Battaglia et aL., 1998). Current radiation-use efficiency is dependent on vapour pressure deficit (VPD), temperature, and light and should decrease as the leaves age and with increased respiration as tree size increases (Mariscal et al, 1999). A decrease in efficiency effectively translates into less structural carbon sequestrated per unit water used or light intercepted. Linder (1985), in a study comparing species across Sweden, New Zealand and Australia showed a linear trend for light interception plotted against AGB production. The study compares pine and Eucalyptus species with deviations from the linearity being attributed to species differences or climatic stresses. A negative intercept for the linear trend in his study was attributed to varying levels of below ground allocation. The quantity of AGB produced per unit of intercepted PAR has been estimated as 0.45 g Mrl for Eucalyptus species in Australia (Beadle, 1997).
The quantity of light absorbed at a gIven LAI and the increase of light absorption with cumulative LAI is dependent on the canopy extinction coefficient (k) (Gazarini et al., 1990).
Intercepted PAR can be estimated by multiplying PAR with an interception fraction calculated using the equation after (Linder, 1985):
1= 10
x
e-k x LAI Equation 13Where1is the flux density of radiation below the canopy and 10 is the flux density of radiation above the canopy,LAJ is leaf area index and k is the light extinction coefficient.
Dividing I by la gives the fraction of non-intercepted light below the forest canopy. This light fraction as recorded by the PCA is plotted as an exponential function (Figure 3.3). The light fraction recorded by the PCA assumes no scattering of light by the foliage (Ll-COR, 1992).
Equation 14 shows the exponential function (derived from Table 16) describing the reduction in below canopy light with increasing LA!. The constant in the equation may describe a 35% light interception by non-foliar portions of the tree canopy and the -k is the canopy extinction coefficient as in Equation 13. Plotting PA! and the fraction of non-intercepted light below the forest canopy give a constant of I (data not shown). It is therefore assumed that the constant in the LAI-based relationship is related to branch and stem attributes. The value of k calculated in this study is approximately 0.50 (Table 16) and is similar to that found in many other studies that estimate utilisable light in relation to LA! (Landsberg and Waring, 1997).
FractlOn. 0f 'non-mtercepted I' IIglt= .0651 xe .-0495xLAI Equation 14
Table 16: Nonlinear regression
predicting extinction coefficient from leaf area index and fraction of light below the tree canopy.
d.f. m.s.
Regression 2 0.0789889Fp..-oOOI
Residual 10 0.0002494
Total 12 0.0133727
1 0.835
r-
SE 0.0158
Constant 0.651Il'r<O.OOI
-k _0.49511''''0.001
0.18 0.16 , 0.14 c: 0.12 .2(j
~ 0.10
~,g> 0.08
~Q.'" 0.06 ~
'"
.~
~C: 0.040 0.02 ~
z
0.00 ,.
2.0 2.5 3.0 3.5
LAI
4.0 4.5
•
5.0 5.5
Figure 3.3 LAI plotted against non-intercepted light fraction.
Annual gridded estimates of radiation obtained from gridded estimates (Schulze, 1997) were converted into PAR by multiplying solar radiation by a factor of 0.45 (Newbould, 1967). With PAR as la, the extinction coefficient (Table 16) and allometrically derived LAI are used to calculated1using Equation 13. Intercepted PAR was calculated as the difference between I and
10. Above-ground productivity per unit of intercepted light energy was calculated by dividing annual AGB by iPAR.
Annual total PAR, iPAR and above-ground productivity expressed in energy units (gMr') of intercepted light energy are shown in Table 17. Ttis assumed that the LAl measured in the 5-11 year age group is a fair reflection of the average LAl during the rotation. This assumption may not hold for the 3-5 year old stands as the early development of LAI and growth rate are unknown, and can be strongly influenced by growth resource availability (du Toit and Dovey, 2004), hence it is excluded from Table 17.
The relationship between intercepted PAR and AGB is shown in Figure 3.4 to demonstrate the deviation of the 3 - 5 year age class from the older classes. A simple line is plotted through the 5 - 11 year old tree data to visually demonstrate the closeness to a linear relationship. Deviation from the line in young stands (Fig 3.4) may be due to higher initial LAl or faster growth as more leaves may be exposed to light, or soil water levels may have been higher early in the rotation. No regression was performed as this is not based on incremental growth data in relation to LAl or actual solar radiation. Solar radiation estimates were not adjusted for cloud cover effects as cloud cover was unknown.
Table 17: A1U1Ual PAR, iPAR, and AGB production per unit light energy for tlu'ee sites in the KwaZulu-Natal Midlands.
AGB/iPA
Site Annual PAR iPAR R
Age (Gl m-2yr-I) (Gl m-2yr-I) (g Mrl)
Bloemendal 5.8 3.60 2.83 0.56
\1istley 5.s 3.61 2.95 0.56
;;eeleMtH 7.3 3.71 3.33 0.77
310emenda 7.8 3.60 2.80 0.46
vlistley 7.0 3.61 2.94 0.53
;eeleMtH 8.2 3.61 3.28 0.69
310emendal 9.8 3.62 2.68 0.41
viistley 11.2 3.50 2.88 0.50
ieeleMtH 11.2 3.49 3.23 0.68
30 X
~ X3-5
'='... 28
~
1.
5.7'='lll:::. 24
.c:
26: .7-9.9-11•
l:0
.'
U:::l 22 X
, , •
'0 20,
,
...
0,
Q. 18 X
,
'0
,
l::::l~~> 1614;
" • ". "t"
".
0 12 .
.c
"
« 10
•
2.5 2.7 2.9 3.1 3.3
intercepted iPAR (GJ.m-2.y(1)
Figure 3.4 The relationship between AGB production and intercepted radiation (iPAR) for stands of different age.
I
3.5,
Closure of the pinnules dw-ing water stress or high evaporative demand may also have caused a more negative intercept than that of Linder, (1985) in Figure 3.4. Stomata are found on both the adaxial and abaxial sides of the leaves, with the abaxial stomata remaining exposed after the pinnules have closed. It is unknown as to whether or not the stomata remain open after the pinnules close, allowing for continued transpiration. If they do remain open, then the leaves will continue to photosynthesise, but perhaps at a lower rate per unit total surface area as a result of a lower level of light interception. Pinnule closw-e may serve to partially reduce water loss, along with increased leaf heat loss by increased air movement. The interception of light by non-foliar pOltions of the tree canopy may also affect the relationship in Table 17, Figure 3.4 as these relate all intercepted light into AGB. Stem and perhaps branch area in relation to LAI changes with age and may contribute to different levels of light interception at various ages. This may need to be explored further with defoliated canopies in order to futther the understanding of this Issue.