Leaf number, water stress and carbon nutrition effects on
poplar leaf growth
J.P. Gaudillère
Laboratoire de Chimie Biologique et de
Photophysiologie,
INRA, 78850 Thivervaf, FranceIntroduction
The final area of a leaf is described
by
thenumber and the mean size of
epidermal
cells. Water stress,
nitrogen, light
andtemperature
have been demonstrated to affectfoliage development.
These factors act at different levels in the process of leafproduction.
The mainsusceptible physio- logical
mechanisms are cell division and differentiation in the apex and in the un-folding
leaf. It has been shown that carbon nutrition has a verysignificant
effect onthe duration of the cell division
phase.
When the leaf
begins
toexport carbon,
cell divisions end
(Dale
andMilthorpe, 1983).
Plant-water relations alsoplay a major
rolethrough
their effects on cellexpansion (Lockhart, 1965).
Cell divisionsare less affected
by
water stress(Clough
and
Milthorpe, 1975).
These
experiments
weredesigned
tospecify
wateravailability
and carbon nutri-* Dr..oco.ont !r/rI,.r::H:!CO’ cfor7nn i1o DhBlcoin.llV’lia B/ónót!lo 1fBICA A
tion effects on the number and the mean
size of
epidermal poplar
leaves.Materials and Methods
Plant production
The
experiments
were carried out onPopulus
euramericana var. 1214.
Cuttings
grew on peatand mineral nutrients were
supplied
as low solubility fertilizer (Osmocote, 18/11/10}. Plantswere watered
daily
with tap water. Growthconditions were: metal halide lamps (200
pmol M
- 2 - S - 1
); light period:
16 h, day andnight
tem- perature and humidity: 20°C and 70%.Anatomical measurements
Areas of attached leaves were measured by the
dot
grid
method (Bouchon, 1975). The numberand the size of
epidermal
cells were measuredon
prints
(Austin et al., 1982) at the final har- vest.RP131 ii1 4n Pnnt-eip-in-KAqvp Fr"n""
*
Present address: Station de Physiologie Vtsg!tale, INRA, BP 131, 33140 Pont-de-la-Maye, France.
Experimental
conditionsLong-term
stress was obtained by lowering the dailywatering
to 10% of the amount of tran-spired
water at thebeginning
of the experiment.After 20 d, full watering was
applied. C0 2
enrichment was carried out in a gas-tight
growth
chamber with the same othergrowing
conditions.
CO!
concentration was controlled and monitoredwith
an IRGA and the hourlyC0
2
uptake
or evolution wascomputed
(La-pierre
et al., 1983).Results
Leaf number
(LN)
effectUnder our
growth conditions,
the final leafarea of successive leaves on control
plants
increased until a constant value of200
(± 10) cm 2 , (Fig. 1 The
number ofcells per leaf increased with LN
(Fig. 2).
The mean cell size was lower in preex-
isting
leaves than in the bud(Fig.
3; LN1-7). Newly
initiated leaves hadinitially large epidermal
cells(LN 8-25).
Then aregular
decrease of the mean leaf areawas observed.
Water stress effect
Reduction of
watering provoked
waterstress. Leaf
growth
was inhibited untilrewatering.
The final leaf size was verysignificantly
decreased(Fig. 1).
Thisdecrease was
entirely explained by
themean size of the cells
(Fig. 3).
The num-ber of cells per leaf
(Fig. 2)
washardly
affected
by
the treatment. With the rewa-tering,
cell divisions andexpansion
startedagain. Newly expanded
leavespresented
the same anatomical characteristics as
the control
(Figs.
2 and3).
Effect of carbon nutrition
A 40% increase in the
photosynthetic C0 2
assimilation rate of the trees was
produ-
ced
by C0 2
enrichment(600 p H - 1 )
of theair. The final leaf area was
highly
stim-ulated
by
this treatment(Fig. 1 The
maingrowth component
affectedby
the in-crease of the carbon nutrition was the number of cells per
leaf,
which doubled(Fig. 2).
The mean size of the cells was not affected(Fig. 3).
Discussion
Water stress and carbon nutrition
signi-
ficantly change
the final area of leaves. It is well known that the watersupply
reduces the
expansive growth
of leaf cells(Cleland, 1971)
and that water stress reduces the final size of theepidermal
cells
(McCree
andDavies, 1974).
Thepresent
results show thatpoplar
leaves follow the samepattern.
The mean size of the cells wasmainly
reducedby
waterstress. Carbon nutrition
specifically changes
the number ofepidermal
cells. Itcan be concluded that cell divisions in an
expanding
leaf are limitedby
the carbonsupply.
In controlplants,
aregular
in-crease in the size of the
newly
initiatedleaves was observed. It could be ex-
plained by
an increased carbonavailability
with the number of leaves of the young tree. The
importance
of thisC0 2
effectcould be
justified, considering
that itincreases the
photosynthetic activity
ofgrowing
leaves.Autotrophy
ofexpanding
leaves in
poplar
has been demonstrated(Larson et aL, 1980)
and carbon nutrition stimulation could beparticularly significant by stimulating autotrophic
carbonsupply
in young leaves.
Acknowledgments
This work was supported by the CNRS
contract:
Physiologie
de la Croissance et duD6veloppement
des V6g6taux ligneux.References
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