Energy requirement for foliage construction depends on tree size inyoung Picea abies trees |
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Authors: | Ülo Niinemets |
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Institution: | LS Pflanzen?kologie II, BIT?K, Universit?t Bayreuth, Postfach 10 12 51, D-95440 Bayreuth, Germany, DE
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Abstract: | The relationship between stand biomass production, and tree age and size is generally a curve with a maximum. To understand
why wood production decreases in the final stages of stand development, the influence of increasing tree size on foliage chemical
composition and substrate requirement for foliage construction in terms of glucose CC, g glucose (g dry mass) –
1] was investigated in the evergreen conifer Picea abies (L.) Karst. Because it was already known that irradiance affects both foliage morphology and chemistry in this species, and
it was expected that the foliage in large overstory trees would intercept on average more light than that in saplings in understory,
irradiance was measured in the sampling locations and included in the statistical models. CC of needles increased with increasing
total tree height (TH) and was independent of relative irradiance. A major reason for increasing CC with increasing TH was
a greater proportion of carbon-rich lignin in the needles in large trees. However, lignin did not fully account for the observed
changes in CC, and it was necessary to assume that certain other carbon-rich secondary metabolites such as terpenes also accumulate
in the foliage of large trees. Enhanced requirements for needle mechanical strength as evidenced by greater lignin concentrations
in large trees were attributed to increased water limitations with increasing tree height. Because water relations may also
control the sink capacities for assimilate usage, apart from the mechanical requirements, they may provide an explanation
for the accumulation of other energetically expensive compounds in the needles as well. Biomass partitioning within the shoot
was another foliar parameter modified in response to increasing tree size. The proportion of shoot axes, which serve to provide
needles with mechanical support and to supply them with water, decreased with increasing TH. This may limit water availability
in the needles, and/or manifest a lower water requirement of the needles containing proportionally more supporting and storage
substances, and consequently, less physiologically active compounds such as proteins. Probably the same factors which caused
CC of the needles to depend on TH, were also responsible for greater CC of the shoot axes in larger trees. These results collectively
suggest that increasingly more adverse water relations with increasing tree size may provide a mechanistic explanation for
the decline in foliar biomass and its functional activity during stand ageing.
Received: 9 April 1996 / Accepted: 14 January 1997 |
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Keywords: | Carbon concentration Construction cost Irradiance Lignin concentration Needle morphology |
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