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PE EIE[R-Rg RESEARCH ON - HJ Andrews Experimental Forest

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ported little photosynthate basipetally in contrast<br />

with the preferential translocation of<br />

photosynthate toward the stem by 2-year-ol d<br />

needles may be explained by the ability of a<br />

sink such as the stem to mobilize photosynthate<br />

falling off with distance in a n<br />

approximately logarithmic manner .<br />

These studies give clues to the relationships<br />

between CO 2 assimilation in the leaves an d<br />

the formation and differentiation of new cells<br />

in apical and cambial growth . These relationships<br />

are not simple, and care must be take n<br />

to avoid oversimplified connections . For<br />

example, net assimilation rates can give an<br />

index of amounts of photosynthate availabl e<br />

for growth only if position and age of th e<br />

foliage studied are taken into account (Ros s<br />

1972). With caution, however, net assimilation<br />

rates taken in different parts of th e<br />

crown can be used to estimate daily mean<br />

assimilation and the total photosynthate production,<br />

on the basis of the whorls o f<br />

branches of different heights (Woodman<br />

1968, 1971) .<br />

In the Coniferous Biome studies, effort s<br />

must be made to bring together net assimilation,<br />

translocation, and terminal and lateral<br />

growth into a workable model . It may be<br />

necessary in such efforts to take into accoun t<br />

growth inhibitors and promoters in such a<br />

model, since Lavender and Hermann (1970 )<br />

have pointed out the importance of thes e<br />

substances in their studies of light and photo -<br />

period effects in Douglas-fir . Although there<br />

is information to build on with respect to re d<br />

pine and Douglas-fir (see above), further wor k<br />

will be necessary with Douglas-fir to utiliz e<br />

the information effectively . Clearly, specifi c<br />

studies of translocation and growth in wester n<br />

hemlock, ponderosa pine, and a true fir ar e<br />

needed .<br />

Mineral Nutrition<br />

A number of books cover the general field<br />

of mineral nutrition and specifically that of<br />

forest tree species (Baule and Fricker 1967 ,<br />

Bengtson 1968, Epstein 1972) . The aspects of<br />

particular concern in the Coniferous Biom e<br />

are mineral uptake and cycling, and the role<br />

of mineral elements in metabolic processes .<br />

Mineral cycling studies in second-growt h<br />

Douglas-fir forests have been carried out for<br />

over a decade (Grier and Cole 1972) . Mineral<br />

cycling is important in the nutrition of th e<br />

trees since limited nutrient capital is available .<br />

These studies need to be extended to th e<br />

other species of special importance in th e<br />

Coniferous Biome program .<br />

The importance of nitrogen in chlorophyll<br />

production and the close correlation between<br />

chlorophyll contents and CO 2 uptake were<br />

pointed out by Keller and Wehrmann (1967 )<br />

with reference to Picea abies and Pinus sylvestris<br />

. The widespread occurrence of nitrogen<br />

deficiency in Douglas-fir in western Washington<br />

and Oregon (Gessel et at . 1965) give s<br />

reason for careful attention to this element i n<br />

studies of coastal Douglas-fir in the Coniferous<br />

Biome . Also it will be wise to make a<br />

sufficiently broad spectrum of mineral analyses<br />

to detect low levels of other element s<br />

which might be limiting photosynthesis or<br />

growth .<br />

The foregoing indicates that a large back -<br />

ground of information on terrestrial production<br />

processes in woody plants is already<br />

available. With respect to Douglas-fir, the<br />

species currently receiving most emphasis i n<br />

the Coniferous Biome, the background i s<br />

appreciable and useful, but often lacking i n<br />

specific data needed for the construction of<br />

mathematical models . These deficiencies must<br />

be made up for adequate modeling of processes<br />

in this species. With other species of<br />

special concern in the Biome-western hemlock,<br />

ponderosa pine, and true firs-muc h<br />

more background information as well as<br />

specific data is needed in order to develo p<br />

effective production models .<br />

A cknowledgments<br />

The work reported in this paper was sup -<br />

ported in part by the University of Washingto n<br />

and in part by National Science Foundatio n<br />

Grant No. GB-20963 to the Coniferous Fores t<br />

Biome, U .S. Analysis of Ecosystems, Inter -<br />

national Biological Program . This is Contribution<br />

No . 37 to the Coniferous <strong>Forest</strong> Biome .<br />

221

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