Table Of ContentOTHER TITLES IN THIS SERIES
1. Plant Modification for More Efficient Water Use
by J.F. Stone (Editor)
1975 χ + 320 pp.
2. Tree Ecology and Preservation
by A. Bernatzky
1978 viii + 358 pp.
3. Cycling of Mineral Nutrients in Agricultural Ecosystems
by M.J. Frissel (Editor)
1978 viii + 356 pp.
4. Interactions Between Non-Pathogenic Soil Microorganisms and Plants
by Y.R. Dommergues and S.V. Krupa (Editors)
1978 xii + 476 pp.
5. Ecology of Root Pathogens
by S.V. Krupa and Y.R. Dommergues (Editors)
1979 χ + 282 pp.
6. Soil Disinfestation
by D. Mulder (Editor)
1979 xiv + 368 pp.
7. Management of Semi-arid Ecosystems
by B.H. Walker (Editor)
1979 χ + 398 pp.
8. Management and Analysis of Biological Populations
by B.S.Goh
1980 χ + 288 pp.
9. Comparison of Forest Water and Energy Exchange Models
by S. Halldin (Editor)
1979 xi + 258 pp.
10. Alternative Methods of Agriculture
by R. Boeringa (Editor)
1980 vi + 200 pp.
11. Wetlands of Bottomland Hardwood Forest
by J.R. Clark and J. Benforado (Editors)
1981 xviii + 402 pp.
12. Plant Production and Management under Drought Conditions
by J.F. Stone and W.O. Willis (Editors)
1983 vii + 390 pp.
13. Evapotranspiration from Plant Communities
by M.L. Sharma (Editor)
1984 vii + 344 pp.
14. Forest Amelioration
by O. Riedl and D. Zachar
1984 viii + 624 pp.
15. Floodplain Forest Ecosystem
by M. Penka, M. Vyskot, E. Klimo and F. Vasicek
Part I. Before Water Management Measures
1985 xii + 468 pp.
Part II. After Water Management Measures
in preparation
16. The Origin and Domestication of Cultivated Plants
by C. Barigozzi (Editor)
1986 vi + 218 pp.
17. Soil Microbial Associations
by V. Vancura and F. Kunc (Editors)
1987
in preparation
18. Tree Crop Physiology
by M.R. Sethuraj and A.S. Raghavendra (Editors)
1987 xii + 361 pp.
DEVELOPMENTS IN AGRICULTURAL
AND MANAGED - FOREST ECOLOGY 18
tree crop
physiology
Edited by
M.R.SETHURAJ
Rubber Research Institute of India, Kottayam 686 009, Kerala, India
and
A.S. RAGHAVENDRA
School of Life Sciences, University of Hyderabad, Hyderabad 500 134, India
ELSEVIER
Amsterdam — Oxford — New York — Tokyo 1987
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© Elsevier Science Publishers B.V., 1987
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ν
PREFACE
Tree crops pose methodological problems for the physiologists, mainly
because of their large size, variable canopy architecture and the perennial
nature. Measurements made on a single leaf or group of leaves, representing
one or different strata are used to describe the physiology of the whole
tree canopy. Often the estimates of tree biomass are only derived from
the measurements of stem-girth and tree-height. The relevance of such
data to the actual situation on tree is, of course, limited. Nevertheless,
the practical problems involved in making realistic measurement of any
physiological parameter at whole-tree level are to be admitted. Therefore,
a logical compromise is sought between practical limitations and the required
accuracy.
Some of the tree crops are vegetatively propagated and the stock-scion
interaction interferes with the desired homogeneity of plant material during
experimental observations. Many tree crops are cross pollinated and the
degree of genetic differences among the population introduces high variability
in the research data. Even when genetically uniform materials are vegeta-
tively propagated, the heterogeneity of soils, because of the large area
required for experimentation, vitiates results in spite of the best statistical
designs. The fluctuations in light climate and other micrometeorological
parameters within a canopy make the attempts to measure photosynthesis
at canopy level, quite laborious. The obstacles on the path of research
in tree physiology created several g'aps in our knowledge. The substantial
amount of loss in biomass due to respiration has not been accurately esti-
mated in tropical tree crops. Similarly, trees could not easily be subjected
to the classical growth analysis or assimilate partioning (including harvest
index estimates) because of their forbidding volume, strong periodicity
in growth and marked remoteness of sink from source (in terms of space
VI
as well as time). These methodological problems have been tackled to
certain extent by individual scientists by adopting innovative modifications
in the methodologies to suit the peculiar situations posed by the tree crops.
Many mathematical principles have been tried and attempts for modelling,
based on representative data, have met with some measure of success.
The scientists who assembled at an international symposium "TREE-
PHYSINDIA-82", held at Rubber Research Institute of India, Kottayam
discussed the special problems encountered by tree physiologists. The plenary
session of this symposium recommended that a publication be brought
out to present information on methodology adopted for various physiological
studies in tree crops.
We made an attempt to present a spectrum of research efforts directed
to understand the trees. The present book therefore contains reviews on
general physiology as well as detailed information on certain selected tree
crops. The contributed articles in the first part, would provide an insight
into different approaches to study the physiology of tree crops, with an
emphasis on principles of methodology. The chapters of the second part,
offer case-histories of physiological investigations on selected economically
important tropical tree crops. Considering the diverse subjects covered,
and the wide variation in the emphasis given to different physiological
aspects among the crops, uniformity of presentation could not be adhered
to.
We would be glad, if this publication forms a source of knowledge
and stimulates further work in the difficult and challenging field of tree
physiology.
M.R. SETHURAJ
January, 1987 A.S. RAGHAVENDRA
VII
ACKNOWLEDGEMENTS
This book is the result of a recommendation from the plenary session
of "TREEPHYSINDIA-82", an international symposium held at Rubber Re-
search institute of India, Kottayam. The editors wish to express their thanks
to the organisers of the symposium for entrusting this task to them. We
wish to acknowledge the permission granted by the Chairman, Rubber
Board and specially thank him for encouragement. One of the editors
(Dr. A.S. Raghavendra) has .since then moved to the University of Hyderabad.
We are grateful to Professor Bh. Krishnamurthy, Vice-Chancellor and
Professor K. Subba Rao, Dean of Life Sciences, University of Hyderabad
for their support and encouragement.
We wish to thank all the authors who have contributed articles in
response to our request. The secretarial assistance of Shri R. Babu is ack-
nowledged. The help of Ms. D. Sudha Sundari and Ms. T. Vani in proof
reading and in the preparation of subject index is gratefully appreciated.
We thank Mr. Anil S. Raina of Instant Business Forms for doing a commen-
dable job in typing this manuscript.
We thank Drs. Jan 3.M. Velterop and Elsevier Science Publishers
for an active association in this venture.
M.R. SETHURAJ
A.S. RAGHAVENDRA
XI
CONTRIBUTORS
The figures in parentheses refer to the number of contributed chapter.
M.C. ANDERSON (4), Commonwealth Scientific and Industrial Research
Organisation, Division of Water and Land Resources, P.O. Box 1666,
Canberra A.C.T. 2601, Australia.
D. BALASIMHA (13), Central Plantation Crops Research Institute, Regional
Station, Vittal 574 243, Karnataka, India.
D.N. BARUA (11), Adviser, Tocklai Experimental Station, Jorhat 785 008,
Assam, India.
E.F. BRUENIG (7), Chair for World Forestry, University of Hamburg and
Institute for World Forestry and Ecology, Hamburg, West Germany.
J. CATSKY (2), Institute of Experimental Botany, Czechoslovak Academy
of Sciences, Flemingovo n.2, CS-16 0 00 Praha 6, Czechoslovakia.
R.H.V. CORLEY (8), Unifield T.C. Limited,Unit 1, St. Martins Ways, Industrial
Estate, Cambridge Road, Bedford MK42 OLG, England, U.K.
F.E. ECKARDT (1), Institute of Plant Ecology, University of Copenhagen,
Oster Farimagsgade 2D, 1353 Copenhagen, Denmark.
N.G. HEGDE (15), The Bharatiya Agro-Industries Foundation, 'Kamdhenu',
Senapati Bapat Road, Pune 411 016, India.
T. KIRA (6), Lake Biwa Research Institute, Otsu, Shiga 520, Japan.
H. MULLERSTAEL (7), Institute for World Forestry and Ecology, Federal
Research Centre for Forestry and Forest Products, Hamburg, West
Germany.
C.A. PRIESTLEY (5), East Mailing Research Station, Maidstone, Kent ME19
6B3, England, U.K.
A.S. RAGHAVENDRA (10), School of Life Sciences, University of Hyderabad,
Hyderabad 500 134, India.
V. RAJAGOPAL (9), Central Plantation Crops Research Station, Kudlu,
Kasaragod 670 124, Kerala, India.
A. RAMADASAN (9), National Research Centre for Spices, Post Box No.1701,
Marikunnu, Calicut 673 012, Kerala, India.
P.K. RAMAIAH (12), Central Coffee Research Institute, Coffee Research
Station 577 117, Karnataka, India.
Z. SESTAK (2,3), Institute of Experimental Botany, Czechoslovak Academy
of Sciences, Flemingovo n.2, CS-16 0 00 Praha 6, Czechoslovakia.
XII
M.R. SETHURA3 (10), Rubber Research Institute of India, Rubber Board,
Kottayam 686 009, Kerala, India.
R.N. SINGH (14), Shyama Sadan, Nand Nagar Colony, I.T.I. Road, Varanasi
411 016, India.
G.R. SQUIRE (8), Unifield T.C. Limited, Unit 1, St. Martins Ways, Industrial
Estate, Cambridge Road, Bedford MK 42 OLG, England, U.K.
D. VENKATARAMANAN (12), Central Coffee Research Institute, Coffee
Research Station 577 117, Karnataka, India.
3. ZIMA (3), Institute of Experimental Botany, Czechoslovak Academy
of Sciences, Flemingovo n.2, CS- 160 00 Praha 6, Czechoslovakia.
3
CHAPTER 1
THE CONTROLLED-ENVIRONMENT PLANT-CHAMBER TECHNIQUE FOR
C0 -EXCHANGE MEASUREMENTS IN TREE AND FOREST RESEARCH
2
F.E. ECKARDT
Institute of Plant Ecology, University of Copenhagen, Oster Farimagsgade 2 D,
1353 Copenhagen, Denmark
ABSTRACT
CO^-exchange is an extremely sensitive indicator of the physiological
behaviour of trees, in relation to the environment, complementary to the
much slower tree ring growth. Among the different methods, employed to
study CO^-exchange, the controlled-environment plant-chamber technique
permits to establish a link between laboratory research and micrometeorologi-
cal observations. This approach implies a thorough understanding of the
metabolism of the O^-exchange as well as of the interaction between
the plant and its environment at different levels of integration in the eco-
system. These matters are considered along with the problem of chosing
the right experimental material. Various control systems employed in plant
chambers are described. Finally, the discussion focusses on deduction of
plant behaviour from chamber data, through a detailed study of not only
various environmental parameters, but also of the architecture of the entire
tree stand.
INTRODUCTION
The complex physiological response of an individual tree to various
environmental factors depend, not only on the plant taxon involved, physical
and biological interactions within the ecosystem of a given climate and
geology, but also on the time and space horizons. Hence it is important to
select the biological and environmental variables at different levels of in-
tegration in the ecosystem, and to study these processes, for each species.
The first approach to such research is to establish a carbon budget
6
of the system. It implies the simultaneous study of CC^-exchange and growth
combined with some modeling. From the budget, it is possible to know the
amount of atmospheric carbon absorbed by the above ground biomass and
the extent of carbon returned to the atmosphere from soil surface. The
carbon budget will also reveal the quantity of the carbon transferred to
the roots (Fig. 1).
Fig. 1. Carbon budget data for Salix glauca stand during a full growing
season. The carbon in carbon dioxide taken up from (+) or given off (-)
to the atmosphere is given on the left. On the right is shown how the carbon
taken up is used for the growth and development of leaves and shoots (-),
or is translocated to the roots (+).
CC^-exchange is measured at leaf or branch level and related to
various weather conditions. Although often combined with growth analysis,
CO^-exchange measurements per se, represent a considerable interest since
they link and predict plant response to specific environmental changes.
Such measurements thus, can partly substitute long-term forest research
programmes (Mooney et al., 1980; Golley, 1983).
PLANT METABOLISM
CO^-exchange
Under optimal water conditions and normal CO£ concentration of
the air, photosynthesis increases proportionally with photon flux density at
low densities but levels off at higher densities. Similarly, under optimal
