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AUSTRALIAN JOURNAL OF
PLANT PHYSIOLOGY
Volume 24, 1997
© CSIRO Australia 1997

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 Aust. J. Plant Physiol., 1997, 24, 381–387

Stomatal Responses to Leaf-to-Air Vapour Pressure Deficit

in Sahelian Species

João P. MarocoAC, João S. PereiraB and M. Manuela ChavesB

A
Department of Botany, Washington State University, Pullman, WA 99164-4238, USA.
B
Instituto Superior de Agronomia, Tapada da Ajuda, Lisboa, Portugal.
C
Corresponding author; e-mail: jpmaroco@wsu.edu

Abstract. Stomatal response to leaf-to-air vapour pressure deficit (LAVPD) was studied in the annual
arid zone C4 grasses Schoenefeldia gracilis, Dactyloctenium aegyptium and Eragrostis tremula and in
the C3 species, convolvulus, Ipomoea pes-tigridis and Ipomoea vagans. Stomatal responses to LAVPD
were consistent with the drought survival strategies adopted by the different species. In drought resistant
species (S. gracilis, I. vagans and I. pes-tigridis) stomatal conductance showed a negative response to
increasing LAVPD whereas, in drought escaping species (D. aegyptium and E. tremula), stomatal
conductance was independent of LAVPD. These observations suggest that resistance to drought was
associated with stomatal closure as LAVPD increased, thus reducing the negative effect of a higher
evaporative demand on water use efficiency, whereas in drought escaping species stomata showed no
response to increasing evaporative demand in the atmosphere.

Keywords: C4, C3, stomatal conductance, leaf-to-air vapour pressure deficit, drought, water-use efficiency

Introduction deficit (LAVPD)*(Grantz 1990). Although it was suggested

Semiarid climates are characterised by rainfall irregular
both in distribution and intensity. It is not unusual in semiarid
climates for more than 10% of the annual rainfall (generally
less than 600 mm/year) to fall in a few hours and for the next
shower to occur only after a few weeks. Between rains there
is, consequently, frequent atmospheric drought (i.e. a high
evaporative demand in the atmosphere) even when soil water
reserves are considerable. Plant survival under these
environmental conditions has evolved mainly through two
different ‘strategies’: (1) drought escape: completion of the
life-cycle after a significant rainfall and before the onset of
the dry season (generally in less than 100 days); and (2)
drought resistance: restricting water losses during droughts
(for a review, see e.g. Turner 1986).
Restricted water loss arises mainly from anatomical
morphological adaptations such as highly cutinised
epidermis or leaf rolling (for reviews, see e.g. Begg 1980;
Berry and Downton 1982) and through stomatal function
(for reviews, see e.g. Ludlow 1980; Schulze 1986). Stomatal
closure under low-humidity atmosphere or soil is an early
and quick response to drought (Cowan 1977; Ludlow 1980;
Schulze 1986). It is accepted that the ambient air humidity
variable sensed by stomata is the leaf-to-air vapour pressure
that peristomatal evaporation mediated this response in a
kind of feedforward response (Raschke 1975; Farquhar
1978; Jarvis and Morison 1981; Sheriff 1982; Dai et al.
1992), it is now accepted that the responses of stomata to
changes in air humidity are in fact the response to changes
in the rate of transpiration (Mott and Parkhurst 1991;
Schulze 1994). It was suggested that localised water deficits
in the mesophyll rather than peristomatal water loss might
be responsible for that response (Schulze 1994), but other
mechanisms may be involved. For example, Kerstiens
(1996) suggested that cuticular transpiration might interact
with such local water deficits in determining the stomatal
response to LAVPD.
In spite of considerable evidence showing that stomata
tend to close with increasing LAVPD in most plant species
(Schulze and Hall 1982; Grantz and Meinzer 1990; Aphalo
and Jarvis 1991), there are apparently some species with
non-responsive stomata (see Ludlow 1980; Sheriff 1982;
Bunce 1988). Indeed, the extent of stomatal responses to
LAVPD shows strong variation dependent on species and
growth conditions, especially air humidity (Kawamitsu et al.
1993), water stress (Tewolde et al. 1993; Schulze 1994),
light (Grantz et al. 1987) and phenological stage (Jones
1992). Non-stomatal responses of photosynthesis to LAVPD

*Abbreviations used: A, net CO2 assimilation rate; A/gs , intrinsic water use efficiency; A/E, instantaneous water use efficiency; Ci, intercellular
CO2 concentration; E, transpiration; gs, stomatal conductance; LAVPD, leaf-to-air vapour pressure deficit; WUE, water-use efficiency.

10.1071/PP96062 0310-7841/97/030381
382
have also been reported (e.g. Sharkey 1984; Bunce 1988).
However, Dai et al. (1992) presented evidence that the
observed results might be artifacts and that the variations in
photosynthesis under varying LAVPD could be accounted
for by changes in stomatal conductance (gs).
Due to its functional significance in intrinsic (A/gs) and
instantaneous (A/E) water use efficiency (Cowan 1977;
Comstock and Ehleringer 1993), stomatal response to
LAVPD has been a topic of current research (Grantz 1990).
As stomata control the diffusion of CO2 and water vapour,
their responses to atmospheric evaporative demand (that is
to leaf-to-air vapour pressure deficit), are fundamental in
controlling the efficiency (and opportunity) of CO2
assimilation regarding water losses. Farquhar et al. (1980)
and Cowan (1982) suggested that stomatal responses to
LAVPD are oriented towards photosynthetic optimisation of
transpiration. However, in semiarid climates, where dry air
is not necessarily accompanied by soil water shortage,
stomata closing at high LAVPD might be ecologically
disadvantageous. If stomata close in response to a dry
atmosphere (i.e. to a high LAVPD, which is especially
frequent in semiarid climates between rainfalls) opportunity
for photosynthesis might be lost when there is still enough
water available in the soil to ensure normal transpiration
rates. This should be particularly detrimental for ‘drought
escaping’ plants that have to fulfill their life-cycle before the
onset of the dry season. On the contrary, for ‘drought
resistant’ plants, stomata closing at high evaporative demand
in the atmosphere could be an efficient mechanism to restrict
water losses and save available soil-water for further growth
(Davies and Pereira 1992).
In this work we studied the stomatal responses to LAVPD
of both drought escaping Dactyloctenium aegyptium (L.)
P. Beauv. and Eragrostis tremula (Lam.) Steud. and drought
resistant Schoenefeldia gracilis Kunth, Ipomoea pes-tigridis
L. and Ipomoea vagans Baker found in the Sahel (Grouzis
1988; Elberse and Breman 1989). The ecological
implications of these responses on water-use efficiency and
drought survival ‘strategies’ will be discussed. The data
presented suggest that stomata which are non-responsive to
LAVPD are associated with drought avoidance ‘strategies’
while stomata responsive to LAVPD are an important
component of drought resistance.
Materials and Methods
Plant Material
The Sahel dominant C4 grasses Schoenefeldia gracilis Kunth,
Dactyloctenium aegyptium (L.) P. Beauv. and Eragrostis tremula
(Lam.) Steud. and the C3 convolvulus Ipomoea vagans Baker and
Ipomoea pes-tigridis L. were obtained from seeds collected in
Dahra-Djollof, Senegalese Sahel (15°238N,15°308W). Seedlings
were grown in a temperature controlled greenhouse in Lisbon
(38°428N, 9°118W) in 12.5 cm diameter pots (7.4 L vol.) filled with
a 1:1 sand-peat mixture and fertilised with 1 kg of commercial
J.P. Maroco et al.
fertiliser per m3 (15:11:13:2 plus Micronutrients, Osmocote, JLV,
Lisbon). Maximum daily natural irradiance in the greenhouse was
502 ± 47.8 mmol m–2 s–1, daily temperature range averaged
20–35°C and relative humidity varied between 15 and 85%. The
plants were watered every 2 days to keep a pre-dawn leaf water
potential of –0.50 ± 0.2 MPa throughout the growing period.
Gas Exchange
Gas exchange measurements were made with a Walz Compact
Mini-Cuvette system with the GK-022 measuring chamber (Heinz
Walz Gmbh, Effeltrich, Germany) on recently fully expanded
leaves. At the time of the measurements plants were entering the
flowering stage. This phenological state was chosen because the
plant responses to drought at this stage are essential to survival (see
e.g. Fischer and Turner 1978; Myers et al. 1992). LAVPD responses
were taken at ambient CO2 ('350 ppm) and at the optimal leaf
temperature for photosynthesis and stomatal aperture previously
determined and shown to be within the natural range of
temperatures of this species habitat. These leaf temperatures were
chosen to avoid factors other than LAVPD that would be limiting
for stomatal aperture and carbon assimilation. For example, if the
leaf temperature was higher than the optimum for photosynthesis,
e.g. in the C3 plants, the decrease in the photosynthetic rate and the
increase in Ci might interact with the stomatal response to LAVPD.
Leaf temperature was measured with a thermocouple and the upper
leaf surface was perpendicularly irradiated with constant saturating
PPFD provided by a ‘L-400 special light fiber optics’ (Heinz Walz
Gmbh, Effeltrich, Germany). LAVPD variations in the cuvette were
obtained by varying air humidity in the inflow measuring gas,
assuming water vapour saturation for intercellular spaces at leaf
temperature. Humidity of the measured gas flow was tracked by
high-precision dew-point mirrors in a CNF-400 bypass flow
control unit (Heinz Walz Gmbh, Effeltrich, Germany). Dew-point
of the inflow measuring gas was kept constant by a flow of dry air
to compensate transpiration. With this system, constant LAVPD
was achieved 3–5 min after changing initial conditions. Net CO2
assimilation (A), transpiration (E) and stomatal conductance to
water vapour (gs) were calculated on-line by the Walz data analysis
program Diagas (ver 2.16) which uses the von Caemmerer and
Farquhar (1981) general gas exchange formulae.
Four measurements were made in different plants of each
species both with sequential increase and decrease of LAVPD.
Readings were taken after steady-state gas exchange rates were
achieved (30 – 40 min after LAVPD change). The curves were
obtained in 2.5 – 3.5 h during daytime (both in the morning and in
the afternoon). Standard exponential, power and polynomial
functions were fitted to observed gs, A, E, A/gs and A/E responses
to LAVPD by least-squares best fits, squared correlation
coefficients and analysis of residues.
Results and Discussion
Typical gas exchange responses to LAVPD for both C3
and C4 plants similar to those observed by e.g. Morison and
Gifford (1983) were observed in I. vagans (Fig. 1a), a
typical ‘drought resistant’ C3 plant (Grouzis 1988). Stomatal
conductance (gs) increased with decreasing LAVPD in a
power type function (explaining 76%*** of total variance).
Stomatal Responses to Leaf-to-Air VPD
The changes in net photosynthesis could be to a large extent
(86%***) accounted for by the variation in gs. Although the
increase in gs by lower LAVPD promoted a higher A, the
increase was less than proportional to the increase in gs,
suggesting that at high conductances the importance of
stomatal limitation to photosynthesis decreased. This might
be explained by an increase in resistance to CO2 diffusion
due to increased collisions with water molecules (Nobel
1991) and by the uncoupling of mesophyll carboxylation
capacity from stomatal conductance, when the stomata are
fully open at very low values of LAVPD. Therefore, as
LAVPD decreased, intrinsic water use efficiency (A/gs)
decreased. The increase in instantaneous water use
efficiency (A/E) as LAVPD decreased was achieved by a
passive reduction in E, due to the reduction in the
evaporating ‘driving force’ and by the increase in CO2
uptake as a result of the opening of stomata.
Gas-exchange responses of I. pes-tigridis to LAVPD were
similar to those observed in I. vagans (Fig. 1b). In this
species stomatal behavior showed, however, a clear
hysteresis with a greater stomatal sensitivity to LAVPD
during the closing phase (increasing LAVPD) than during the
opening phase (decreasing LAVPD), as reported for other
species (Barradas et al 1994).
Assuming that stomatal closure at high LAVPD resulted
from local water stress in the mesophyll cells, possibly not
detectable if xylem water potential were measured (Schulze
1994), this hysteresis might result from one or two
mechanisms. One possibility would be a lag in rehydration
of the guard cell complex after the lowest LAVPD was
reached. This might be more important if the hydraulic
conductance within the leaves is low. Another possibility
would be the persistence of ABA in the guard cell
environment after stomatal closure. It is possible that there
might be a release of ABA from the mesophyll cells
associated with the local water deficits (Slovik and Hartung
1992). The redistribution of ABA would serve to keep the
stomata closed after reaching a low LAVPD rather than
inducing the stomatal response to LAVPD (Kerstiens 1996).
Moreover, Hartung and Slovik (1991) modelled the effects of
increasing LAVPD in ABA concentrations in the guard-cell
environment and concluded that these concentrations may
increase after exposure to dry air and that, if LAVPD is low
enough, it would take some time for [ABA] to return to the
initial values. Even though this would explain the observed
hysteresis in stomatal response to LAVPD, this hypothesis has
not been clearly proven. Differences between species
regarding the hysteresis response might therefore result from
differences in hydraulic properties of leaf tissues or in
sensitivity of the stomata to ABA.
Most of the variation in stomatal conductance for this
‘drought resistant’ species was explained by LAVPD
(94%*** and 98%*** for increasing and decreasing LAVPD
respectively) with an abrupt reduction in gs for LAVPD
greater than 20 Pa/kPa. The variation in A was satisfactorily
383
explained by changes in gs (63%***). Stomatal opening as a
result of LAVPD changing from 25 to 15 Pa/kPa, caused an
abrupt increase in transpiration which was lost as the
evaporation driving force decreased with further reductions
in LAVPD. At high LAVPD, gs decreased more than
proportionally as compared to A, leading to high intrinsic
WUE (A/gs). This reflects an efficient stomatal control of
water losses by this species. As for the instantaneous WUE
(A/E), the highest values were recorded at both high and low
LAVPD. However, while for high LAVPD the high A/E was
due to stomatal control of gas exchange at low LAVPD, the
high A/E arose passively as a result of the low evaporative
demand on transpiration. Therefore, I. pes-tigridis should be
particularly efficient in water use for CO2 assimilation under
the extreme conditions of air humidity (high and low).
Furthermore, the morphological characteristics of the leaves
of this species, namely dense epidermal hairs, help to
prevent water loss from leaf surfaces at high evaporative
demands (Berry and Dowton 1982; Bòlhar-Nordenkampf
and Draxler 1993). The shading of the leaf surface caused by
these morphological characteristics could also help to
prevent other stresses, namely high temperature and high
light, on the photoassimilatory apparatus.
The gas-exchange response to LAVPD of S. gracilis
(Fig. 1c) was also compatible with this species ‘drought
resistance’ behaviour (Grouzis 1988; Elberse and Breman
1989) as discussed above for the C3 convolvulus species.
Stomatal conductance increased exponentially with
decreasing LAVPD (R2=0.93**), with the observed variation
in A being satisfactorily explained by gs responses
(R2=0.59*). However for this species, intrinsic WUE (A/gs)
tends to be constant for low and medium LAVPD. As for the
instantaneous WUE (A/E), the highest values resulted from
the passive reduction of E as LAVPD decreased. The strategy
of S. gracilis under water stress seems to result in water
saving rather than keeping a high instantaneous water use
efficiency.
Both ‘drought avoiding’ species E. tremula and
D. aegyptium (Grouzis 1988; Elberse and Breman 1989)
lacked stomatal sensitivity to LAVPD (Figs 1d and 1e).
Indeed, the best fitted power function for E. tremula and
exponential for D. aegyptium’s gs response to LAVPD
explain less than 10%ns of total variability, showing no effect
of LAVPD in the stomatal function. As suggested by Sheriff
(1982) for Tradescantia virginiana which responds weakly
to LAVPD when the plants are well watered (Schulze 1994),
high hydraulic conductance in the leaves could at least in
part compensate for the evaporation losses from the
mesophyll and prevent the occurrence of localised water
deficits. For the species studied, a high proportion of thin-
wall bulliform cells (for D. aegyptium) and high vascular
density (for E. tremula) (data not shown) could lead to the
high hydraulic conductance of leaves and the absence of
response to LAVPD. Indeed, thin cell walls in opposition to
384
thick cell walls (more suberised cell walls) have higher
hydraulic conductivity according to Sheriff (1982). It makes
sense as it is known that more-suberised cell walls have
fewer plasmodesmatal connections (symplast transport) and
the higher thickness of the cell wall reduces water transport
efficiency (by increasing the length of the apoplastic
pathway). It is possible that, as found in other species, water
stressed leaves show a greater sensitivity to high LAVPD
(Schulze 1994).
Intrinsic WUE of both species (D. aegyptium and
E. tremula) was independent of LAVPD as A also had no
response to this atmospheric variable, as could be expected
if direct, non-stomatal effects of LAVPD on photosynthesis
do not occur (Dai et al. 1992). Increasing instantaneous
WUE with decreasing LAVPD was, thereafter, explained
satisfactorily (more than 57%** for both species) by the
reduction of transpiration due to the decreasing evaporation
driving force as LAVPD decreases (Figs 1d and 1e). The
response of these species to atmospheric drought seems to
rely only on a passive response of transpiration to the
LAVPD. This should not be necessarily an ecological
disadvantage. Stomatal functioning of E. tremula and
D. aegyptium seems oriented for maximising gas exchange
independent of atmospheric dryness. This is compatible with
a full usage of soil-water resources and the fulfillment of the
life cycle after a significant rainfall and before the onset of
soil-drought, and it may be supported by a high growth
potential at early stages of life of these C4 species (Maroco
1995). If stomata closed in response to a high evaporative
demand in the atmosphere, the opportunity for carbon
assimilation would be lost even when the soil reservoir was
able to insure a normal transpiration flow, which is usually
the case between rainfall events. Stomata closing at high
LAVPD could indeed be disadvantageous for completion of
the life-cycle before the real onset of the dry-season.
Although we did not measure non-stomatal transpiration,
it is plausible to assume that this alternative pathway of
water loss might play a role in stomatal responses to high
LAVPD. As suggested by Kerstiens (1996), the degree of
stomatal sensitivity to LAVPD may correlate with cuticle
permeability. Whereas species with higher cuticular water
permeance may have more sensitive stomatal responses to
LAVPD, less permeable cuticules would lead to lower
stomatal sensitivy to the LAVPD (Kerstiens 1996). This may
explain the lack of sensitivity of both D. aegyptium and
E. tremula, where more cutinised epidermal cells were
observed (Maroco 1995), comparatively to the other species
in this study (unpublished observations).
Conclusions
This paper shows two contrasting responses to
atmospheric humidity (sensed as LAVPD): whereas some
species showed the more usual pattern of decrease in
J.P. Maroco et al.
stomatal conductance in dry air, others were non-sensitive to
LAVPD changes. We may speculate that these stomatal
responses to atmospheric humidity are compatible with the
drought survival strategies postulated for the Sahelian
species studied. For the ‘drought resistant’ I. vagans, I. pes-
tigridis and S. gracilis, stomatal closure in response to
increasing atmospheric drought reduced soil water usage,
preventing rapid exhaustion of soil moisture reserves.
Besides other morphological adaptations (such as dense
epidermal hairs or leaf rolling) this stomatal behaviour
might restrict soil-water usage enabling a sort of
feedforward response to the occurrence of drought.
Although opportunities for carbon uptake may be lost at
times when soil moisture might provide enough water for
‘normal’ transpiration flow, this strategy would insure plant
survival until the next rain and, therefore, the fulfillment of
the longer life-cycle of these plants.
The strategy of ‘drought escaping’ species, E. tremula
and D. aegyptium, which have the C4 type of photosynthesis,
would be to make maximum use of the available soil
moisture in order to maximise their carbon assimilation so
that they can complete their life cycle before the onset of
soil-drought. Even though we have not studied the responses
of stomata to air humidity under soil water stress, we may
speculate that stomatal insensitivity to LAVPD, and stomatal
closing only in response to soil water depletion, could be
hazardous if the next rainfall were to occur too late in the
season. Other characteristics found in these species, like
seed dormancy requiring intense rainfall to leach out the
inhibitory substances (Maroco 1995), might prevent
germination when the rainfall is not enough to insure plant
growth and development. These seed characteristics, as well
as other morphological responses such as leaf rolling, can
compensate for stomatal insensitivity to high LAVPD by
reducing transpiration and preventing lethal dehydration at
times of very high evaporative demand. It must be
emphasised, however, that the stomatal non-responsiveness
to high LAVPD of well watered plants would ensure a
competitive advantage over other species for minerals, light
and water.
Acknowledgments
To Drs T. Nolan, J. Connoly and Agronomist C. Shall for
logistic support in seed collecting and to Dr M. E. Poulson
and two anonymous reviewers for the critical review of the
manuscript. This work was financed by the EU Project,
Contract Nr. TS3-CT93-0202. Financial support to one of us
(JPM) was provided by a scholarship from the ‘Junta
Nacional de Investigação Cientifica e Tecnológica’
(Contract No: BM/3112/92-IE), Lisbon, Portugal.
 (a) Ipomoea vagans (b) Ipomoea pes-tigridis (c) Schoenefeldia gracilis (d) Eragrostis tremula (e) Dactyloctenium aegyptium

Stomatal Responses to Leaf-to-Air VPD

Fig. 1. Stomatal conductance (A), net photosynthesis (B), transpiration (C), intrinsic WUE (A/gs ) (D) and instantaneous WUE (A/E) (E) response to LAVPD. Different symbols refer to
different plants. Open symbols, measurements made with decreasing LAVPD; closed symbols, measurements made with increasing LAVPD. (a) I. vagans measured at 27.7 ± 0.0°C leaf
temperature, 1157 ± 3 mmol m –2s –1 PPFD and pre-dawn leaf y = –0.3 ± 0.1 MPa. Solid line, best fitted power function as described in Material and Methods. (b) I. pes-tigridis measured
at 26.1 ± 0.0°C leaf temperature, 1178 ± 19 mmol m –2s –1 PPFD and predawn leaf y = –0.3 ± 0.1 MPa. Solid line, best 3rd-order polynomial function as described in Material and Methods.
(c) S. gracilis measured at 29.8 ± 0.1°C leaf temperature, 1459 ± 6 mmol m –2s –1 PPFD and predawn leaf y = –0.6 ± 0.2 MPa. Lines, best fitted 2nd-order polynomial function as described
in Material and Methods. (d) E. tremula measured at 34.0 ± 0.1°C leaf temperature, 1182 ± 12 mmol m –2s –1 PPFD and pre-dawn y = –0.5 ± 0.1 MPa. Solid line, best fitted power
function as described in Material and Methods. (e) D. aegyptium measured at 30.2 ± 0.1°C leaf temperature, 1477 ± 6 mmol m –2s –1 PPFD and pre-dawn leaf y = –0.4 ± 0.1 MPa. Solid
line, best fitted exponential function as described in Material and Methods.

385
386
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Manuscript received 13 June 1996, accepted 27 February 1997