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Journal of Experimental Botany, Vol. 55, No.

407,
Water-Saving Agriculture Special Issue, pp. 24272436, November 2004
doi:10.1093/jxb/erh213 Advance Access publication 30 July, 2004

Irrigation scheduling: advantages and pitfalls


of plant-based methods

Hamlyn G. Jones*
Plant Research Unit, Division of Environmental and Applied Biology, School of Life Sciences,
University of Dundee at SCRI, Invergowrie, Dundee DD2 5DA, UK

Received 12 November 2003; Accepted 27 May 2004

Abstract into the realms of use, or at most the refinement, of existing

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practical applications. Nevertheless, in recent years there
This paper reviews the various methods available for
has been a wide range of proposed novel approaches to
irrigation scheduling, contrasting traditional water-
irrigation scheduling which have not yet been widely
balance and soil moisture-based approaches with
adopted; many of these are based on sensing the plant
those based on sensing of the plant response to water
response to water deficits rather than sensing the soil
decits. The main plant-based methods for irrigation
moisture status directly (Jones, 1990a).
scheduling, including those based on direct or indirect
The increasing worldwide shortages of water and costs of
measurement of plant water status and those based on
irrigation are leading to an emphasis on developing
plant physiological responses to drought, are outlined
methods of irrigation that minimize water use (maximize
and evaluated. Specic plant-based methods include
the water use efficiency). The advent of precision irrigation
the use of dendrometry, fruit gauges, and other tissue
methods such as trickle irrigation has played a major role in
water content sensors, while measurements of growth,
reducing the water required in agricultural and horticultural
sap ow, and stomatal conductance are also outlined.
crops, but has highlighted the need for new methods of
Recent advances, especially in the use of infrared
accurate irrigation scheduling and control. In recent years it
thermometry and thermography for the study of sto-
has become clear that maintenance of a slight plant water
matal conductance changes, are highlighted. The rela-
deficit can improve the partitioning of carbohydrate to
tive suitabilities of different approaches for specic
reproductive structures such as fruit and also control
crop and climatic situations are discussed, with the
aim of indicating the strengths and weaknesses of
excessive vegetative growth (Chalmers et al., 1981), giving
different approaches, and highlighting their suitability
rise to what has been termed by Chalmers et al. (1986) as
over different spatial and temporal scales. The poten-
regulated deficit irrigation (RDI). Achievement of suc-
tial of soil- and plant-based systems for automated
cessful RDI depends on accurate soil moisture or plant
irrigation control using various scheduling techniques
stress sensing, and requires an ability to irrigate little and
is also discussed.
often on demand. A disadvantage of RDI is that it requires
water status to be maintained accurately within a rather
Key words: Dendrometry, sap-ow, stomatal conductance, narrow tolerance; any excess application loses the advan-
thermography, water balance. tage of the regulated deficit and can cost more in terms of
water used, while any under-application can lead to severe
yield or quality losses. An alternative recent innovation to
achieve the same measure of growth control has been the
Introduction
development of partial root-zone drying (PRD), where
Irrigation scheduling has conventionally aimed to achieve irrigation is supplied alternately to different parts of the root
an optimum water supply for productivity, with soil water system (Dry and Loveys, 1998; Stoll et al., 2000b). A
content being maintained close to field capacity. In many potential advantage of this method is that precise irrigation
ways irrigation scheduling can be regarded as a mature control is probably less critical for success than it is for
research field which has moved from innovative science RDI, as plants can always obtain adequate water from the

* Fax: +44 1382 34275. E-mail: h.g.jones@dundee.ac.uk

Journal of Experimental Botany, Vol. 55, No. 407, Society for Experimental Biology 2004; all rights reserved
2428 Jones
well-watered side of the root system and the drying side different crops and different climates, as required in the
primarily provides a signal to modify growth and stomatal water balance calculation, have been reviewed in detail by
aperture (Stoll et al., 2000a). Allen et al. (1999). Although the water balance approach
The range of crops to which RDI and PRD methods have is not very accurate, it has generally been found to be
been applied is increasing all the time, but their greatest sufficiently robust under a wide range of conditions.
successes have been in high-value horticultural and fruit Nevertheless it is subject to the serious problem that errors
crops, usually those where the harvested part of the plant is are cumulative over time. For this reason it is often
its reproductive organ. Applications of such techniques to necessary to recalibrate the calculated water balance at
extensive arable crops are in their infancy, although there intervals by using actual soil measurements, or sometimes
are some exciting preliminary reports (Kang et al., 2000, plant response measurements (as outlined below). Some of
2003). At present it is much less clear whether PRD or RDI the main advantages and disadvantages of the different
would be so valuable for vegetative crops, although irrigation scheduling approaches are outlined in Table 1.
appropriate application can be used to restrict growth, as A potential problem with all soil-water based approaches
is required for high quality in some ornamental crop species is that many features of the plants physiology respond
(RS Harrison-Murray, personal communication). directly to changes in water status in the plant tissues,
The choice of irrigation scheduling method depends to whether in the roots or in other tissues, rather than to
a large degree on the objectives of the irrigator and the changes in the bulk soil water content (or potential). The
irrigation system available. The more sophisticated sched- actual tissue water potential at any time therefore depends

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uling methods generally require higher-precision applica- both on the soil moisture status and on the rate of water flow
tion systems; nevertheless even less sophisticated systems through the plant and the corresponding hydraulic flow
such as flood irrigation scheduling can benefit from resistances between the bulk soil and the appropriate plant
improvements in irrigation scheduling as outlined here. tissues. The plant response to a given amount of soil
The pressures to improve irrigation use efficiency and to moisture therefore varies as a complex function of evap-
use irrigation for precise control of vegetative growth, as in orative demand. As a result it has been suggested (Jones,
RDI, both imply a requirement for increased precision in 1990a) that greater precision in the application of irrigation
irrigation control, maintaining the soil moisture status can potentially be obtained by a third approach, the use of
within fine bands to achieve specific objectives in crop plant stress sensing. For this approach irrigation
management. Such objectives can only be met by precision scheduling decisions are based on plant responses rather
irrigation systems such as trickle irrigation that can apply than on direct measurements of soil water status; some of
precise amounts of water at frequent intervals (often several the possible physiological measurements and responses that
times per day). Effective operation of such systems equally can be used are discussed in the following section.
requires a sensing system that determines irrigation need in
real time or at least at frequent intervals; this rules out large-
scale manual monitoring programmes for such purposes Plant-based methods for irrigation control
and indicates a need for automated monitoring systems.
Introduction
If soil water-based measures are to be replaced by plant-
Basics of irrigation scheduling
based measures it is important to consider what measures
The main methods that are used for irrigation scheduling, or might be most appropriate for irrigation scheduling pur-
that have the potential for development in the near future, poses. Possible measures include direct measurements of
are summarized in Table 1. Irrigation scheduling is con- some aspect of plant water status as well as measurements of
ventionally based either on soil water measurement, a number of plant processes that are known to respond
where the soil moisture status (whether in terms of water sensitively to water deficits. One might expect that a direct
content or water potential) is measured directly to de- measure of plant water status should be the most rigorous
termine the need for irrigation, or on soil water balance and hence the most useful indicator of irrigation require-
calculations, where the soil moisture status is estimated by ment, although the question remains as to where in the plant
calculation using a water balance approach in which the that quantity should be measured. In practice, as has been
change in soil moisture (Dh) over a period is given by the argued strongly by Jones (1990b), most plants exercise
difference between the inputs (irrigation plus precipitation) some measure of autonomous control over their shoot or leaf
and the losses (runoff plus drainage plus evapotranspira- water status, tending to minimize changes in shoot water
tion). Soil moisture measurement techniques have been the status as the soil dries or as evaporative demand increases
subject of many texts and reviews (Smith and Mullins, (Bates and Hall, 1981; Jones, 1983). In the long term, this
2000; Dane and Topp, 2002) and will not be addressed control is achieved through changes in leaf area and root
here. Similarly, the detailed methods for estimating evapo- extension, and in the shorter term through changes in leaf
transpiration and calculation of crop water requirements for angle, stomatal conductance, and hydraulic properties of the
Irrigation scheduling 2429
Table 1. A summary of the main classes of irrigation scheduling approaches, indicating their main advantages and disadvantages
Comments that relate to all methods in a section are not repeated in subsections.
Advantages Disadvantages

I. Soil water measurement


(a) Soil water potential (tensiometers, Easy to apply in practice; can be quite precise; Soil heterogeneity requires many sensors (often
psychrometers, etc.) at least water content measures indicate how expensive) or extensive monitoring programme
much water to apply; many commercial (e.g. neutron probe); selecting position that is
systems available; some sensors (especially representative of the root-zone is difficult;
capacitance and time domain sensors) readily sensors do not generally measure water status
automated at root surface (which depends on evaporative
demand)
(b) Soil water content (gravimetric;
capacitance/TDR; neutron probe)
II. Soil water balance calculations
(Require estimate of evaporation and rainfall) Easy to apply in principle; indicate how much Not as accurate as direct measurement; need
water to apply accurate local estimates of precipitation/runoff;
evapotranspiration estimates require good
estimates of crop coefficients (which depend
on crop development, rooting depth, etc.);
errors are cumulative, so regular recalibration
needed

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III. Plant stress sensing
(Includes both water status measurement and Measures the plant stress response directly; In general, does not indicate how much water
plant response measurement) integrates environmental effects; potentially to apply; calibration required to determine
very sensitive control thresholds; still largely at research/
development stage and little used yet for routine
agronomy (except for thermal sensing in some
situations)
(a) Tissue water status It has often been argued that leaf water status is All measures are subject to homeostatic
the most appropriate measure for many regulation (especially leaf water status), therefore
physiological processes (e.g. photosynthesis), not sensitive (isohydric plants); sensitive to
but this argument is generally erroneous (as it environmental conditions which can lead to
ignores rootshoot signalling) short-term fluctuations greater than treatment
differences
(i) Visible wilting Easy to detect Not precise; yield reduction often occurs before
visible symptoms; hard to automate
(ii) Pressure chamber (w) Widely accepted reference technique; most Slow and labour intensive (therefore expensive,
useful if estimating stem water potential (SWP), especially for predawn measurements);
using either bagged leaves or suckers unsuitable for automation
(iii) Psychrometer (w) Valuable, thermodynamically based measure of Requires sophisticated equipment and high
water status; can be automated level of technical skill, yet still unreliable in
the long term
(iv) Tissue water content (RWC, leaf Changes in tissue water content are easier to Instrumentation generally complex or expensive,
thickness [c- or b-ray thickness measure and automate than water potential so difficult to get adequate replication; water
sensors], fruit or stem diameter) measurements; RWC more directly related to content measures (and diameter changes)
physiological function than is total water subject to same problems as other water status
potential in many cases; commercial measures; leaf thickness sensitivity limited by
micromorphometric sensors available lateral shrinkage
(v) Pressure probe Can measure the pressure component of water Only suitable for experimental or laboratory
potential which is the driving force for xylem systems
flow and much cell function (e.g. growth)
(vi) Xylem cavitation Can be sensitive to increasing water stress Cavitation frequency depends on stress
prehistory; cavitationwater status curve shows
hysteresis, with most cavitations occurring
during drying, so cannot indicate successful
rehydration
(b) Physiological responses Potentially more sensitive than measures of Often require sophisticated or complex
tissue (especially leaf) water status equipment; require calibration to determine
control thresholds
(i) Stomatal conductance Generally a very sensitive response, except in Large leaf-to-leaf variation requires much
some anisohydric species replication for reliable data
Porometer Accurate: the benchmark for research studies Labour intensive so not suitable for commercial
application; not readily automated (though some
attempts have been made)
Thermal sensing Can be used remotely; capable of scaling up to Canopy temperature is affected by
large areas of crop (especially with imaging); environmental conditions as well as by stomatal
imaging effectively averages many leaves; aperture, so needs calibration (e.g. using wet
simple thermometers cheap and portable; well and dry reference surfaces)
suited for monitoring purposes
2430 Jones
Table 1. Continued
Advantages Disadvantages

Sap-flow sensors Sensitive Only indirectly estimates changes in


conductance, as flow is also very dependent
on atmospheric conditions; requires complex
instrumentation and technical expertise; needs
calibration for each tree and for definition of
irrigation control thresholds
(ii) Growth rate Probably the most sensitive indicator of water Instrumentation delicate and generally
deficit stress expensive

transport system. In extreme cases, plants with good endog- Reduction in tissue (MPa) required to affect process
enous control systems maintain a stable leaf water status Process affected 0 1 2
over a wide range of evaporative demand or soil water
Cell growth
supplies; these plants are termed isohydric (Stocker, 1956), Wall synthesis
and include especially plants such as cowpea, maize, and Protein synthesis
poplar (Bates and Hall, 1981; Tardieu and Simonneau, Protochlorophyllide
Nitrate synthase
1998). This is by contrast with those species such as ABA accumulation

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sunflower or barley which appear to have less effective Cytokinin concentration
Stomatal opening
control of leaf water status and have been termed anisohy- Photosynthesis
dric. In practice the distinctions between isohydric and Respiration
anisohydric behaviour are often not clear-cut; even different Proline accumulation
Sugar accumulation
cultivars of grapevine have been shown to have contrasting
hydraulic behaviours (Schultz, 2003). Fig. 1. Generalized sensitivities of plant processes to water deficits
The choice of which plant-based measure to use depends (modified with permission from Hsiao, 1973).
on their relative sensitivity to water deficits. The definition
of sensitivity, however, is somewhat problematic. The can also be misleading, because with slowly developing
relative sensitivities of different physiological processes stress the plant adapts by decreasing leaf area; as a result
were reviewed in some detail by Hsiao (1973), who stomatal conductance and photosynthesis rate per unit leaf
identified cell growth as being most sensitive to tissue area may remain fairly stable as soil dries (Moriana and
water deficits, closely followed by wall and protein syn- Fereres, 2002). Nevertheless, over shorter time-scales it still
thesis, all of which could respond to water deficits of less appears that stomata are a particularly sensitive early
than 0.1 MPa (Fig. 1). Hsiao reported that stomatal closure indicator of water deficits.
was only rarely affected when tissue water potential fell by In principle, water status is not ideal as a measure of
0.20.5 MPa, with decreases of 1.0 MPa or more being water deficit as it is already subject to some physiological
required for stomatal closure in many cases. Although control, and indeed, as has been outlined above, leaf water
photosynthesis was classified as moderately sensitive by potential generally shows some homeostasis. Nevertheless,
Hsiao, largely as a result of its dependence on stomatal changes in water status somewhere in the plant system are
aperture, some component processes such as electron assumed to be a prerequisite for any physiological adapta-
transport are now known to be particularly insensitive tion or other response. All that a homeostatic system can do
(Massacci and Jones, 1990). It is now believed that Hsiaos is to minimize, not eliminate, the changes in water status;
(1973) classification is somewhat misleading, and under- indeed for a feedback system of stomatal control it is not
estimates the true sensitivity of the stomata, as it is based on theoretically possible for such a system to stably eliminate
observed responses to leaf water potential alone and changes in shoot water status if that is the variable that
ignores the internal rootshoot signalling that is now actually controls the stomata (Jones, 1990b; Franks et al.,
known to play a major part in controlling stomatal aperture 1997).
(Davies and Zhang, 1991). In general, the use of any plant-based or similar indicator
The error arising from a reliance on leaf water status is for irrigation scheduling requires the definition of reference
readily apparent when one considers that many plants or threshold values, beyond which irrigation is necessary
operate when optimally watered with the leaf water (Table 1). Such reference values are commonly determined
potential at around 2 MPa, yet the stomata may close as for plants growing under non-limiting soil water supply
the soil dries by only a few tens of Pa, with little change in (Fereres and Goldhamer, 2003), but obtaining extensive
leaf water potential (Bates and Hall, 1981). A further information on the behaviour of these reference values as
consideration is that any attempt to relate stomatal aperture environmental conditions change is an important stage in
to leaf water potential in a long-term drought experiment the development and validation of such methods. Another
Irrigation scheduling 2431
general limitation to plant-based methods is that they do not largely equilibrate with wsoil by dawn). Unfortunately this is
usually give information on how much irrigation to apply often found to be rather insensitive to variation in soil
at any time, only whether or not irrigation is needed. moisture content (Garnier and Berger, 1987). Further, this
is not very convenient for irrigation scheduling as routine
Plant water status measurements predawn are expensive to obtain, and at best
Perhaps the first approach to the use of the plant itself as an can only be obtained daily. As yet another alternative, Jones
indicator of irrigation need, and one that is still frequently (1983) suggested the indirect estimation of an effective soil
adopted today, was to base irrigation on visible wilting. water potential at the root surface of transpiring plants
Unfortunately, by the time wilting is apparent a substantial based on measurements of leaf water potential and stomatal
proportion of potential yield may already have been lost conductance during the day, and argued that this should
(Slatyer, 1967). More rigorous and more sensitive measures have significant advantages over predawn measurements.
of plant water status are therefore required. Although Such an approach has been successfully tested by Lorenzo-
relative water content (RWC) (Barrs, 1968) is a widely Minguez et al. (1985).
used measure of water status that does not require sophis- None of the above plant-based methods are well adapted
ticated equipment, it is often argued that water potential, for automation of irrigation scheduling or control because
especially of the leaves (wleaf) is a more rigorous and more of the difficulties of measurement of any of the variables
generally applicable measure of plant water status (Slatyer, discussed. Although it may be possible to use automated
1967; Jones, 1990b). In spite of this, RWC has the stem or leaf psychrometers (Dixon and Tyree, 1984), these

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advantage that it can be more closely related to cell turgor, instruments are notoriously unreliable. In conclusion, it is
which is the process directly driving cell expansion, than it apparent from the above discussion that the favoured way
is to the total water potential (Jones, 1990b). to use plant water status is actually as an indicator of soil
The fact that plant water status, and especially leaf water water status; this negates many of the advantages of
status, is usually controlled to some extent by means of selecting a plant-based measure! Indeed soil water potential
stomatal closure or other regulatory mechanisms, argues can be measured directly, thus avoiding the need for any
against the use of such measures, especially in strongly plant-based measurement, although it is worth noting that
isohydric species. A further problem with the use of leaf this does not necessarily give a good measure of the
water status as an indicator of irrigation need was pointed effective water potential at the root surface during active
out by Jones (1990b), who noted that even though there was transpiration (Jones, 1983).
often homeostasis of leaf water potential between different Several indirect methods for measuring or monitoring
soil moisture regimes, rapid temporal fluctuations are often water status have been developed as alternatives to direct
observed as a function of environmental conditions (such as measurement. The general behaviour of a number of such
passing clouds). This makes the interpretation of leaf water methods have been compared by McBurney (1992) and
potential as an indicator of irrigation-need doubly unsatis- Selles and Berger (1990). In general, these indirect methods
factory. Nevertheless, in spite of the concerns with the use suffer from the same disadvantages as do the direct
of leaf water status that have been outlined above, it has measurements of leaf water status, but in certain circum-
been reported that leaf water potential can, when corrected stances have been developed into commercial systems.
for diurnal and environmental variation, provide a sensitive Some of these approaches are reviewed below.
index for irrigation control (Peretz et al., 1984).
Leaf thickness: A number of instruments are available for
As a partial solution to the variability of leaf water status,
the routine monitoring of leaf thickness, which is known to
various workers have proposed that a more useful and more
decrease as turgidity decreases. Approaches include direct
robust indicator of water status is the xylem water potential
measurement using linear displacement transducers (e.g.
or stem water potential (SWP, measured by using a pressure
LVDTs [Burquez, 1987; Malone, 1993] or capacitance
chamber on leaves enclosed in darkened plastic bags for
sensors [McBurney, 1992]) or through measurements of
some time before measurement and allowed to equilibrate
leaf superficial density using b-ray attenuation (Jones,
with the xylem water potential; McCutchan and Shackel,
1973). Unfortunately, leaf thickness is frequently even less
1992). As a more stable measure of water status, others
sensitive to changes in water status than is leaf water
have even recommended that measurements should be
content because, especially with younger leaves, a fraction
made on pre-equilibrated leaves from root suckers (Jones,
of leaf shrinkage is often in the plane of the leaves rather
1990a; Simonneau and Habib, 1991). These methods are
than in the direction of the sensor (Jones, 1973).
thought to be preferable largely because they approach
more closely the soil water status than does the value of leaf Stem and fruit diameter: Stem and fruit diameters fluctuate
water potential, although as a result they therefore miss out diurnally in response to changes in water content, and so
on the potential advantages of plant-based methods. suffer from many of the same disadvantages as other water
Perhaps an even better estimator of the soil water status measures. Nevertheless, the diurnal dynamics of
potential is the predawn leaf water potential (as wleaf should changes in diameter, especially of fruits, have been used to
2432 Jones
derive rather more sensitive indicators of irrigation need, Although the changes in transpiration rate that sap flow
where the magnitude of daily shrinkage has been used to indicates are largely determined by changes in stomatal
indicate water status, and comparisons of diameters at the aperture, transpiration is also influenced by other environ-
same time on succeeding days give a measure of growth mental conditions such as humidity. Therefore changes in
rate (Huguet et al., 1992; Li and Huguet, 1990; Jones, sap flow can occur without changes in stomatal opening.
1985). Although changes in growth rate provide a particu- Even though rates of sap flow may vary markedly between
larly sensitive measure of plant water stress, such daily trees as a result of differences in tree size and exposure, the
measurements are not particularly useful for the control of general patterns of change in response to both environ-
high-frequency irrigation systems. Nevertheless, several mental conditions and to water status are similar (Eastham
workers have achieved promising results for low-frequency and Gray, 1998). Appropriate sap-flow rates to use as
irrigation scheduling by the use of maximum daily shrink- control thresholds may be derived by means of regular
age (MDS). For example, Fereres and Goldhamer (2003) calibration measurements, especially for larger trees. Al-
showed that MDS was a more promising approach for ternatively, it is at least feasible in principle to derive an
automated irrigation scheduling than was the use of stem irrigation scheduling algorithm that is based on an analysis
water potential for almond trees, while differences in of the diurnal patterns of sap flow, with midday reductions
maximum trunk diameter were also found to be particularly being indicative of developing water deficits (though of
useful in olive (Moriana and Fereres, 2002). The use of course diurnal fluctuations in environmental conditions can
such dendrometry or micromorphometric techniques has mimic such changes). Another potential problem with sap

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been developed into a number of successful commercial flow for precision control is that it tends to lag behind
irrigation scheduling systems (e.g. Pepista 4000, Delta changes in transpiration rate owing to the hydraulic
International, Montfavet, France); these are usually applied capacitance of the stem and other plant tissues (Wronski
to the study of stem diameter changes. Selles and Berger et al., 1985).
(1990) reported that variations in trunk diameter or stem It follows that, although sap-flow measurement is well
water potential were more sensitive as indicators of irriga- adapted for automated recording and hence potentially
tion need than was the variation in fruit diameter. This was automated control of irrigation systems, it can be a little
probably a result of the poor hydraulic connection between difficult to determine the correct control points for any crop.
fruit tissue and the conducting xylem. There is currently
much interest in evaluating such techniques for irrigation Xylem cavitation
scheduling, with a number of relevant papers presented at It is generally accepted (Steudle, 2001) that water in the
recent meetings (e.g. the International Society for Horticul- xylem vessels of transpiring plants is under tension; as
tural Science 4th International Symposium on Irrigation of water deficits increase, this tension is thought to increase to
Horticultural Crops, 15 September 2003, Davis, CA, USA such an extent that the water columns can fracture, or
[as yet unpublished], and Kang et al., 2003). cavitate. Such cavitation events lead to the explosive
formation of a bubble, initially containing water vapour.
c-ray attenuation: A related approach to the study of
These cavitation events can be detected acoustically in the
changes in stem water content was the use of c-ray
audio- (Milburn, 1979) or ultrasonic-frequencies (Tyree
attenuation (Brough et al., 1986). Although this was shown
and Dixon, 1983), and the resulting embolisms may restrict
to be very sensitive, safety considerations and cost have
water flow through the stem. Substantial evidence, though
largely limited the further application of this approach.
largely circumstantial, now indicates that the ultrasonic
acoustic emissions (AEs) detected as plants become
Sap ow stressed do indeed indicate cavitation events and that AE
The development of reliable heat pulse and energy balance rates can be used as an indicator of plant stress (Tyree and
thermal sensors for sap-flow measurement in the stems of Sperry, 1989). Nevertheless, there remain many uncertain-
plants (Granier, 1987; Cohen et al., 1981; Cermak and ties as it seems that at least a proportion of the AEs detected
Kucera, 1981) has opened up an alternative approach to as woody tissues dry out may not be related to xylem
irrigation scheduling based on measurements of sap-flow embolisms. For example, the large numbers observed by
rates. Because sap-flow rates are expected to be sensitive to Sandford and Grace (1985) as coniferous stems dried out
water deficits and especially to stomatal closure, many were substantially in excess of the number of conducting
workers have tested the use of sap-flow measurement for tracheids present, thus suggesting a major contribution to
irrigation scheduling and control in a diverse range of observed AEs by the non-conducting fibres (Jones and
crops, including grapevine (Eastham and Gray, 1998; Pena, 1986). Although the measurement of AEs has proved
Ginestar et al., 1998a, b), fruit and olive trees (Ameglio to be a powerful tool for the study of hydraulic architecture
et al., 1998; Fernandez et al., 2001; Giorio and Giorio, in plants, there has been little progress in adapting this
2003; Remorini and Massai, 2003) and even greenhouse measure as an indicator for irrigation scheduling. Note,
crops (Ehret et al., 2001). however, the recent report by Yang et al. (2003), who
Irrigation scheduling 2433
implemented a control algorithm based on the association (CWSI) by Idso and colleagues (Idso et al., 1981; Jackson
of AE rate with transpiration rate for the precision irrigation et al., 1981), where CWSI was obtained from the canopy
of tomato. It is likely that the main reasons for the lack of temperature (Tcanopy) according to
uptake include the fact that the relationship between the
CWSI = Tcanopy  Tnws =Tdry  Tnws 1
number of AEs and water status changes with successive
cycles of stress, and the fact that cavitation events are where Tnws is a so-called non-water-stressed baseline tem-
mostly observed during the drying phase, not during perature for the crop in question at the same atmospheric
rewetting, and so cannot provide an indicator of when vapour pressure deficit, and Tdry is an independently derived
irrigation has been sufficient to replenish the soil water temperature of a non-transpiring reference crop (Fig. 2). In
supply. this approach all temperatures are expressed as differences
from air temperature so that standard relationships for Tdry
Stomatal conductance and thermal sensing and Tnws can be used. Although this approach was found to
As outlined above, it appears that changes in stomatal be useful in the clear arid climate of Arizona where the
conductance are particularly sensitive to developing water method was developed, it has proved to be less useful in
deficits in many plants and therefore potentially provide more humid or cloudy climates where the signal-to-noise
a good indicator of irrigation need in many species. It is in ratio is somewhat smaller (see Fig. 2; Hipps et al., 1985;
this area that most effort has been concentrated on the Jones, 1999). In spite of its deficiencies, there has been
development of practical, plant-based irrigation scheduling widespread use of infrared thermometry as a tool in

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approaches. Although stomatal conductance can be mea- irrigation scheduling in many, especially arid, situations
sured accurately using widely available diffusion porome- (Jackson, 1982; Stockle and Dugas, 1992; Martin et al.,
ters, measurements are labour-intensive and unsuitable for 1994), especially with the development of trapezoidal
automation. The recognition that leaf temperature tends methods involving the combination of temperature data
to increase as plants are droughted and stomata close with a visible/near infrared vegetation index (Moran et al.,
(Raschke, 1960) led to a major effort in the 1970s and 1994).
1980s to develop thermal sensing methods, based on the In order to improve the precision of the approach in more
newly developed infrared thermometers, for the detection humid or low-radiation environments, Jones (1999) in-
of plant stress (see reviews by Jackson, 1982; Jones and troduced the approach of using physical dry and wet
Leinonen, 2003; Jones, 2004). reference surfaces to replace the notional Tdry and Tnws
An early method of accounting for the rapid short-term required for equation 1. A number of recent papers have
variation in leaf temperature as radiation and wind speed shown that this approach can give reliable and sensitive
vary in the field was to refer leaf temperatures to air tem- indications of stomatal closure (Diaz-Espejo and Verhoef,
perature and to integrate these differences (e.g. the Stress 2002; Jones et al., 2002; Leinonen and Jones, 2004) and
Degree Day measure of Jackson et al., 1977); significant hence has the potential to be used for irrigation scheduling.
elevation of canopy temperature above air temperature was The most important recent advances in the application of
indicative of stomatal closure and water deficit stress. The thermal sensing for plant stress detection and irrigation
method was transformed into a more practical approach scheduling, however, have been provided by the introduc-
following the introduction of the crop water stress index tion of thermal imagery (Jones, 1999, 2004; Jones et al.,

Fig. 2. (A) Illustration of the calculation of Idsos Crop Water Stress Index: CWSI=(TcanopyTnws)/(TdryTnws), showing the dependence of Tnws ( )
and Tdry (- - -) on air vapour pressure deficit (vpd, kPa). (B) Illustration of the effect of a given experimental noise (for example resulting from
measurement errors and variations in irradiance), indicated by the double-headed arrow, showing that the signal-to-noise ratio decreases markedly as the
vpd decreases from levels found in hot and arid/semi-arid climates to values typical in humid or maritime climates.
2434 Jones
2002), although their expense has meant that such systems automated irrigation controller based on thermal sensing of
have yet to be widely used. plant stress. Similar approaches have been applied in the
In addition to the use of the absolute temperature rise as field: for example, Evans et al. (2001) and Sadler et al.
stomata close, it has also been proposed that use may be (2002) mounted an array of 26 infrared thermometers
made of the fact that the variance of leaf temperature (IRTs) on a centre pivot irrigation system which they used
increases as stomata close (Fuchs, 1990). Indeed, this may to monitor irrigation efficiency, but had not developed the
be a more sensitive indicator of stomatal closure than is system to a stage where it could be used for fully automated
the temperature rise (Jones, 2004). Again, the introduction control. Colaizzi et al. (2003) have tested another system
of thermal cameras now makes the wider use of such that includes thermal sensing of canopy temperature on
approaches feasible, especially when combined with a large linear move irrigator (where the irrigator moves
automated image analysis. across the field). In another approach to the use of canopy
temperature that makes use of the thermal kinetic win-
dow, Upchurch et al. (1990) and Mahan et al. (2000) have
Automation
developed what they call a biologically identified optimal
The most widespread use of automated irrigation scheduling temperature interactive console for the control of trickle
systems is in the intensive horticultural, and especially the and other irrigation systems based on canopy temperature
protected cropping, sector. In general, the automated sys- measurements. In this direct control system, irrigation is
tems in common use are based on simple automated timer applied as canopy temperature exceeds a crop-specific

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operation, or in some cases the signal is provided by soil optimum. The development of thermal infrared imaging
moisture sensors. For timer-based operation many systems methods of irrigation control will be aided by the recent
simply aim to provide excess water to runoff at intervals (e.g. development of automated image analysis systems for
flood-beds or capillary matting systems), although some at extraction of the temperatures of leaf surfaces from thermal
least attempt to limit water application by only applying images, including shaded and sunlit leaves, soil, and other
enough to replenish evaporative losses (often calculated surfaces (Leinonen and Jones, 2004).
from measured pan evaporation; Allen et al., 1999). Much
greater sophistication is required if an objective is to improve
the overall irrigation water use efficiency or to apply an RDI Conclusions
system. Most of the remaining automated systems currently This review has briefly considered the current state of the
in operation base control on soil moisture sensing; at least art and potential opportunities for use of plant-based stress
this approach has the potential for greater precision and sensing as the basis for irrigation scheduling and control.
improved water use efficiency. The advantages and disadvantages of each of these ap-
Applications of automated plant-based sensing are largely proaches are summarized in Table 1. Although plant-based
in the developmental stage, partly because it is usually sensing has several potential advantages, including a greater
necessary to supplement the plant-stress sensing by addi- relevance to plant functioning than soil-based measures,
tional information (such as evaporative demand). In princi- these have been offset by a number of practical difficulties
ple, with high-frequency on-demand irrigation systems one of implementation that have thus far limited the develop-
could envisage a real-time control system where water ment of commercially successful systems. However, pres-
supply is directly controlled by a feedback controller sures for enhanced water use efficiency and for greater
operated by the stress sensor itself, so that no information precision in irrigation systems are likely to provide a real
on the required irrigation amount is needed. For such an impetus for the development of new precision irrigation
approach care will be necessary to take account of any lags in scheduling systems that take account of the irrigation need
the plant physiological response used for the control signal. of individual plants, and may well involve greater use of
The use of expert systems (Plant et al., 1992), which plant-based sensing systems.
integrate data from several sources, appears to have great
potential for combining inputs from thermal or other crop
response sensors and environmental data for a water budget Acknowledgements
calculation to derive a robust irrigation schedule.
The author is grateful to sponsors of various aspects of the work pre-
Among the various plant-based sensors that have been sented, who include the European Commission (projects: WATER-
incorporated into irrigation control systems are stem di- USE, contract EVKI-200022061, and STRESSIMAGING, Contract
ameter gauges (Huguet et al., 1992), sap-flow sensors HPRN-CT-200200254).
(Schmidt and Exarchou, 2000) and acoustic emission
sensors (Yang et al., 2003), though there has been most
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