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Effects of N-Fertilization on Instantaneous Carbon Fixation
Ability of Douglas-Fir Foliage: Relative Importance of Leaf Area and Photosynthetic
Rate
Thomas M. Hinckley, Professor, ES, College of Forest
Resources, University of Washington
Zuo Shen and Holly Barnard, Graduate Students
College of Forest Resources, University of Washington
It is well understood that trees respond to nitrogen fertilization by
increasing leaf area and photosynthetic rates. We wanted to investigate
how this nitrogen-mediated response might lead to increases in productivity.
Physiological interactions occur between light environment, leaf area, leaf
nitrogen and photosynthetic capacity and we have done research to explore
these interactions in the context of productivity. The objective of our
research was to examine the relative contribution of increased leaf area
and increased photosynthetic rates on leaf carbon balance caused by nitrogen
fertilization in different canopy positions and among different leaf age
classes. In the top part of the tree crowns, light is not a limiting factor.
The amount of nitrogen available to the leaves is a decisive factor in photosynthesis.
On the other hand, at the bottom of the canopy, because more nitrogen is
allocated to the higher light environments, both light and nitrogen are
limiting. So increasing the amount of nitrogen may or may not increase the
photosynthetic capacity of leaves depending on which factor is more limiting.
Furthermore, fertilization tends to shift the leaf area toward upper canopy
so as to increase the mutual shading in the lower canopy. However if the
foliage in the lower crown is more nitrogen limited, then increasing nitrogen
to these leaves will increase the photosynthetic capacity and partly counterbalance
the negative effect of mutual shading. We are expecting that the amount
of carbon production through nitrogen fertilization contributed by increased
photosynthetic rates would increase from top to bottom of the crown.
BACKGROUND
A 15-year-old Douglas-fir plantation in Snoqualmie Tree Farm was chosen
as the study site. The stand was planted in 1985 with 2-0 seedlings after
a clear-cut on a 40% slope. Measured in the March of 1997, average tree
heights were 11.6 and 11.0 m and DBHs, were 13.7 and 15.7 cm in unfertilized
and fertilized plots, respectively. One stand was fertilized in March of
1993 and again in 1997 with 224 kg N / ha of urea, the other served as a
control. Two trees from each plot were selected for leaf area measurements
by using a systematic branch sampling technique. A model was constructed
to predict leaf area from branch diameter and its relative position within
a trees crown for each treatment. To calibrate the equations, one
big tree in each plot with similar DBH was selected and a modified stratified-clip
sampling technique was used to calibrate the model developed by the systematic
sampling method. Light saturated photosynthesis was also measured in April
of 1998 using a Li-Cor 6200 on several different age classes of foliage
from the 3rd, 5th and 7th whorl which corresponded to the upper, middle,
and lower crown positions. A branchs instantaneous carbon fixation
ability was calculated as the sum of the products of light saturated photosynthetic
rate and leaf area for each age class of foliage on a branch. Then increased
carbon fixation production through N fertilization was partitioned into
two parts: carbon production due to increased photosynthetic rate and carbon
production due to increased foliage area.
RESULTS
The results showed that N fertilization increased leaf area and net photosynthesis
over all canopy positions. Compared to the control, fertilized trees have
120%, 92%, and 100% more leaf area for a given nodal branch in upper, middle,
and lower crown positions respectively (Figure 1).
Figure 1
Light saturated photosynthetic rates of fertilized trees were also 29%,
97%, and 96% higher than the corresponding branches of unfertilized trees
(Figure 2).
Figure 2
In the upper crown position, 85% of the increased carbon production of
foliage by N fertilization was through increased foliage area, while in
the middle and lower crown positions, it was 65% and 68% respectively (Figure
3).
Figure 3
The ssremainder was through increased rates of net photosynthesis. In
terms of current year foliage, more than 90% of the increased carbon fixation
ability was contributed by increased leaf area and less than 10% was contributed
by increased photosynthetic rates and the relative contribution of increased
photosynthetic rates increased as one goes from top to the bottom of the
crown (Figure 4).
Figure 4
Also it appears that the amount of contribution by increased photosynthetic
rates increased from one-year old needles until 3-year-old needles, then
declined for needles in the upper and middle crown positions. There was
no apparent pattern over age for the needles in lower crown positions.
CONCLUSION
We found that fertilized Douglas-fir trees have much more leaf area than
unfertilized trees. Light-saturated rates of photosynthesis increased in
fertilized trees, and this did not have much of a pattern with leaf age.
We feel that the larger productivity of the fertilized trees is mostly due
to their increased leaf area, and secondarily due to their increased photosynthesis.
There is a strong interaction of light and nitrogen. Big trees benefit most
from fertilization.
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