Irrigation scheduling: advantages and pitfalls of plant-based methods

Abstract

RESEARCH PAPER

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

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

Received 12 November 2003; Accepted 27 May 2004


Abstract 

 
This paper reviews the various methods available for irrigation scheduling, contrasting traditional water-balance and soil moisture-based approaches with those based on sensing of the plant response to water deficits. The main plant-based methods for irrigation scheduling, including those based on direct or indirect measurement of plant water status and those based on plant physiological responses to drought, are outlined and evaluated. Specific plant-based methods include the use of dendrometry, fruit gauges, and other tissue water content sensors, while measurements of growth, sap flow, and stomatal conductance are also outlined. Recent advances, especially in the use of infrared thermometry and thermography for the study of stomatal conductance changes, are highlighted. The relative suitabilities of different approaches for specific crop and climatic situations are discussed, with the aim of indicating the strengths and weaknesses of different approaches, and highlighting their suitability over different spatial and temporal scales. The potential of soil- and plant-based systems for automated irrigation control using various scheduling techniques is also discussed.

Key words: Dendrometry, sap-flow, stomatal conductance, thermography, water balance

Source: Journal of Experimental Botany 2004 55(407):2427-2436


Introduction

 

Irrigation scheduling has conventionally aimed to achieve an optimum water supply for productivity, with soil water content being maintained close to field capacity. In many ways irrigation scheduling can be regarded as a mature research field which has moved from innovative science into the realms of use, or at most the refinement, of existing practical applications. Nevertheless, in recent years there has been a wide range of proposed novel approaches to irrigation scheduling which have not yet been widely adopted; many of these are based on sensing the plant response to water deficits rather than sensing the soil moisture status directly (Jones, 1990aGo).

The increasing worldwide shortages of water and costs of irrigation are leading to an emphasis on developing methods of irrigation that minimize water use (maximize the water use efficiency). The advent of precision irrigation methods such as trickle irrigation has played a major role in reducing the water required in agricultural and horticultural crops, but has highlighted the need for new methods of accurate irrigation scheduling and control. In recent years it has become clear that maintenance of a slight plant water deficit can improve the partitioning of carbohydrate to reproductive structures such as fruit and also control excessive vegetative growth (Chalmers et al., 1981Go), giving rise to what has been termed by Chalmers et al. (1986)Go as ‘regulated deficit irrigation’ (RDI). Achievement of successful RDI depends on accurate soil moisture or plant ‘stress’ sensing, and requires an ability to irrigate ‘little and often’ on demand. A disadvantage of RDI is that it requires water status to be maintained accurately within a rather narrow tolerance; any excess application loses the advantage 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 development of partial root-zone drying (PRD), where irrigation is supplied alternately to different parts of the root system (Dry and Loveys, 1998Go; Stoll et al., 2000bGo). A potential advantage of this method is that precise irrigation control is probably less critical for success than it is for RDI, as plants can always obtain adequate water from the well-watered side of the root system and the drying side primarily provides a signal to modify growth and stomatal aperture (Stoll et al., 2000aGo).

The range of crops to which RDI and PRD methods have been applied is increasing all the time, but their greatest successes have been in high-value horticultural and fruit crops, usually those where the harvested part of the plant is its reproductive organ. Applications of such techniques to extensive arable crops are in their infancy, although there are some exciting preliminary reports (Kang et al., 2000Go, 2003Go). At present it is much less clear whether PRD or RDI would be so valuable for vegetative crops, although appropriate application can be used to restrict growth, as is required for high quality in some ornamental crop species (RS Harrison-Murray, personal communication).

The choice of irrigation scheduling method depends to a large degree on the objectives of the irrigator and the irrigation system available. The more sophisticated scheduling methods generally require higher-precision application systems; nevertheless even less sophisticated systems such as flood irrigation scheduling can benefit from improvements in irrigation scheduling as outlined here. The pressures to improve irrigation use efficiency and to use irrigation for precise control of vegetative growth, as in RDI, both imply a requirement for increased precision in irrigation control, maintaining the soil moisture status within fine bands to achieve specific objectives in crop management. Such objectives can only be met by precision irrigation systems such as trickle irrigation that can apply precise amounts of water at frequent intervals (often several times per day). Effective operation of such systems equally 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 and indicates a need for automated monitoring systems.


Basics of irrigation scheduling

 

The main methods that are used for irrigation scheduling, or that have the potential for development in the near future, are summarized in Table 1. Irrigation scheduling is conventionally based either on ‘soil water measurement’, where the soil moisture status (whether in terms of water content or water potential) is measured directly to determine the need for irrigation, or on ‘soil water balance calculations’, where the soil moisture status is estimated by calculation using a water balance approach in which the change in soil moisture ({Delta}{theta}) over a period is given by the difference between the inputs (irrigation plus precipitation) and the losses (runoff plus drainage plus evapotranspiration). Soil moisture measurement techniques have been the subject of many texts and reviews (Smith and Mullins, 2000Go; Dane and Topp, 2002Go) and will not be addressed here. Similarly, the detailed methods for estimating evapotranspiration and calculation of crop water requirements for different crops and different climates, as required in the water balance calculation, have been reviewed in detail by Allen et al. (1999)Go. Although the water balance approach is not very accurate, it has generally been found to be sufficiently robust under a wide range of conditions. Nevertheless it is subject to the serious problem that errors are cumulative over time. For this reason it is often necessary to recalibrate the calculated water balance at intervals by using actual soil measurements, or sometimes plant response measurements (as outlined below). Some of the main advantages and disadvantages of the different irrigation scheduling approaches are outlined in Table 1.

A potential problem with all soil-water based approaches is that many features of the plant's physiology respond directly to changes in water status in the plant tissues, whether in the roots or in other tissues, rather than to changes in the bulk soil water content (or potential). The actual tissue water potential at any time therefore depends both on the soil moisture status and on the rate of water flow through the plant and the corresponding hydraulic flow resistances between the bulk soil and the appropriate plant tissues. The plant response to a given amount of soil moisture therefore varies as a complex function of evaporative demand. As a result it has been suggested (Jones, 1990aGo) that greater precision in the application of irrigation can potentially be obtained by a third approach, the use of ‘plant "stress" sensing’. For this approach irrigation scheduling decisions are based on plant responses rather than on direct measurements of soil water status; some of the possible physiological measurements and responses that can be used are discussed in the following section.



Plant-based methods for irrigation control

 

 
Introduction
If soil water-based measures are to be replaced by plant-based measures it is important to consider what measures might be most appropriate for irrigation scheduling purposes. Possible measures include direct measurements of some aspect of plant water status as well as measurements of a number of plant processes that are known to respond sensitively to water deficits. One might expect that a direct measure of plant water status should be the most rigorous and hence the most useful indicator of irrigation requirement, although the question remains as to where in the plant that quantity should be measured. In practice, as has been argued strongly by Jones (1990b)Go, most plants exercise some measure of autonomous control over their shoot or leaf water status, tending to minimize changes in shoot water status as the soil dries or as evaporative demand increases (Bates and Hall, 1981Go; Jones, 1983Go). In the long term, this control is achieved through changes in leaf area and root extension, and in the shorter term through changes in leaf angle, stomatal conductance, and hydraulic properties of the transport system. In extreme cases, plants with good endogenous control systems maintain a stable leaf water status over a wide range of evaporative demand or soil water supplies; these plants are termed ‘isohydric’ (Stocker, 1956Go), and include especially plants such as cowpea, maize, and poplar (Bates and Hall, 1981Go; Tardieu and Simonneau, 1998Go). This is by contrast with those species such as sunflower or barley which appear to have less effective control of leaf water status and have been termed ‘anisohydric’. In practice the distinctions between isohydric and anisohydric behaviour are often not clear-cut; even different cultivars of grapevine have been shown to have contrasting hydraulic behaviours (Schultz, 2003Go).

The choice of which plant-based measure to use depends on their relative sensitivity to water deficits. The definition of sensitivity, however, is somewhat problematic. The relative sensitivities of different physiological processes were reviewed in some detail by Hsiao (1973)Go, who identified cell growth as being most sensitive to tissue water deficits, closely followed by wall and protein synthesis, all of which could respond to water deficits of less than 0.1 MPa (Fig. 1). Hsiao reported that stomatal closure was only rarely affected when tissue water potential fell by 0.2–0.5 MPa, with decreases of 1.0 MPa or more being required for stomatal closure in many cases. Although photosynthesis was classified as moderately sensitive by Hsiao, largely as a result of its dependence on stomatal aperture, some component processes such as electron transport are now known to be particularly insensitive (Massacci and Jones, 1990Go). It is now believed that Hsiao's (1973)Go classification is somewhat misleading, and underestimates the true sensitivity of the stomata, as it is based on observed responses to leaf water potential alone and ignores the internal root–shoot signalling that is now known to play a major part in controlling stomatal aperture (Davies and Zhang, 1991Go).

The error arising from a reliance on leaf water status is readily apparent when one considers that many plants operate when optimally watered with the leaf water potential at around –2 MPa, yet the stomata may close as the soil dries by only a few tens of Pa, with little change in leaf water potential (Bates and Hall, 1981Go). A further consideration is that any attempt to relate stomatal aperture to leaf water potential in a long-term drought experiment can also be misleading, because with slowly developing stress the plant adapts by decreasing leaf area; as a result stomatal conductance and photosynthesis rate per unit leaf area may remain fairly stable as soil dries (Moriana and Fereres, 2002Go). Nevertheless, over shorter time-scales it still appears that stomata are a particularly sensitive early indicator of water deficits.

In principle, water status is not ideal as a measure of water deficit as it is already subject to some physiological control, and indeed, as has been outlined above, leaf water potential generally shows some homeostasis. Nevertheless, changes in water status somewhere in the plant system are assumed to be a prerequisite for any physiological adaptation or other response. All that a homeostatic system can do is to minimize, not eliminate, the changes in water status; indeed for a feedback system of stomatal control it is not theoretically possible for such a system to stably eliminate changes in shoot water status if that is the variable that actually controls the stomata (Jones, 1990bGo; Franks et al., 1997Go).

In general, the use of any plant-based or similar indicator for irrigation scheduling requires the definition of reference or threshold values, beyond which irrigation is necessary (Table 1). Such reference values are commonly determined for plants growing under non-limiting soil water supply (Fereres and Goldhamer, 2003Go), but obtaining extensive information on the behaviour of these reference values as environmental conditions change is an important stage in the development and validation of such methods. Another general limitation to plant-based methods is that they do not usually give information on ‘how much’ irrigation to apply at any time, only whether or not irrigation is needed.

Plant water status
Perhaps the first approach to the use of the plant itself as an indicator of irrigation need, and one that is still frequently adopted today, was to base irrigation on visible wilting. Unfortunately, by the time wilting is apparent a substantial proportion of potential yield may already have been lost (Slatyer, 1967Go). More rigorous and more sensitive measures of plant water status are therefore required. Although relative water content (RWC) (Barrs, 1968Go) is a widely used measure of water status that does not require sophisticated equipment, it is often argued that water potential, especially of the leaves ({psi}leaf) is a more rigorous and more generally applicable measure of plant water status (Slatyer, 1967Go; Jones, 1990bGo). In spite of this, RWC has the advantage that it can be more closely related to cell turgor, which is the process directly driving cell expansion, than it is to the total water potential (Jones, 1990bGo).

The fact that plant water status, and especially leaf water status, is usually controlled to some extent by means of stomatal closure or other regulatory mechanisms, argues against the use of such measures, especially in strongly isohydric species. A further problem with the use of leaf water status as an indicator of irrigation need was pointed out by Jones (1990b)Go, who noted that even though there was often homeostasis of leaf water potential between different soil moisture regimes, rapid temporal fluctuations are often observed as a function of environmental conditions (such as passing clouds). This makes the interpretation of leaf water potential as an indicator of irrigation-need doubly unsatisfactory. Nevertheless, in spite of the concerns with the use of leaf water status that have been outlined above, it has been reported that leaf water potential can, when corrected for diurnal and environmental variation, provide a sensitive index for irrigation control (Peretz et al., 1984Go).

As a partial solution to the variability of leaf water status, various workers have proposed that a more useful and more robust indicator of water status is the xylem water potential or stem water potential (SWP, measured by using a pressure chamber on leaves enclosed in darkened plastic bags for some time before measurement and allowed to equilibrate with the xylem water potential; McCutchan and Shackel, 1992Go). As a more stable measure of water status, others have even recommended that measurements should be made on pre-equilibrated leaves from root suckers (Jones, 1990aGo; Simonneau and Habib, 1991Go). These methods are thought to be preferable largely because they approach more closely the soil water status than does the value of leaf water potential, although as a result they therefore miss out on the potential advantages of plant-based methods.

Perhaps an even better estimator of the soil water potential is the predawn leaf water potential (as {psi}leaf should largely equilibrate with {psi}soil by dawn). Unfortunately this is often found to be rather insensitive to variation in soil moisture content (Garnier and Berger, 1987Go). Further, this is not very convenient for irrigation scheduling as routine measurements predawn are expensive to obtain, and at best can only be obtained daily. As yet another alternative, Jones (1983)Go suggested the indirect estimation of an effective soil water potential at the root surface of transpiring plants based on measurements of leaf water potential and stomatal conductance during the day, and argued that this should have significant advantages over predawn measurements. Such an approach has been successfully tested by Lorenzo-Minguez et al. (1985)Go.

None of the above plant-based methods are well adapted for automation of irrigation scheduling or control because of the difficulties of measurement of any of the variables discussed. Although it may be possible to use automated stem or leaf psychrometers (Dixon and Tyree, 1984Go), these instruments are notoriously unreliable. In conclusion, it is apparent from the above discussion that the favoured way to use plant water status is actually as an indicator of soil water status; this negates many of the advantages of selecting a plant-based measure! Indeed soil water potential can be measured directly, thus avoiding the need for any plant-based measurement, although it is worth noting that this does not necessarily give a good measure of the effective water potential at the root surface during active transpiration (Jones, 1983Go).

Several indirect methods for measuring or monitoring water status have been developed as alternatives to direct measurement. The general behaviour of a number of such methods have been compared by McBurney (1992)Go and Sellés and Berger (1990)Go. In general, these indirect methods suffer from the same disadvantages as do the direct measurements of leaf water status, but in certain circumstances have been developed into commercial systems. Some of these approaches are reviewed below.

Leaf thickness:
A number of instruments are available for the routine monitoring of leaf thickness, which is known to decrease as turgidity decreases. Approaches include direct measurement using linear displacement transducers (e.g. LVDTs [Burquez, 1987Go; Malone, 1993Go] or capacitance sensors [McBurney, 1992Go]) or through measurements of leaf ‘superficial density’ using ß-ray attenuation (Jones, 1973Go). Unfortunately, leaf thickness is frequently even less sensitive to changes in water status than is leaf water content because, especially with younger leaves, a fraction of leaf shrinkage is often in the plane of the leaves rather than in the direction of the sensor (Jones, 1973Go).

Stem and fruit diameter:
Stem and fruit diameters fluctuate diurnally in response to changes in water content, and so suffer from many of the same disadvantages as other water status measures. Nevertheless, the diurnal dynamics of changes in diameter, especially of fruits, have been used to derive rather more sensitive indicators of irrigation need, where the magnitude of daily shrinkage has been used to indicate water status, and comparisons of diameters at the same time on succeeding days give a measure of growth rate (Huguet et al., 1992Go; Li and Huguet, 1990Go; Jones, 1985Go). Although changes in growth rate provide a particularly sensitive measure of plant water stress, such daily measurements are not particularly useful for the control of high-frequency irrigation systems. Nevertheless, several workers have achieved promising results for low-frequency irrigation scheduling by the use of maximum daily shrinkage (MDS). For example, Fereres and Goldhamer (2003)Go showed that MDS was a more promising approach for automated irrigation scheduling than was the use of stem water potential for almond trees, while differences in maximum trunk diameter were also found to be particularly useful in olive (Moriana and Fereres, 2002Go). The use of such dendrometry or micromorphometric techniques has been developed into a number of successful commercial irrigation scheduling systems (e.g. ‘Pepista 4000’, Delta International, Montfavet, France); these are usually applied to the study of stem diameter changes. Sellés and Berger (1990)Go reported that variations in trunk diameter or stem water potential were more sensitive as indicators of irrigation need than was the variation in fruit diameter. This was probably a result of the poor hydraulic connection between fruit tissue and the conducting xylem. There is currently much interest in evaluating such techniques for irrigation scheduling, with a number of relevant papers presented at recent meetings (e.g. the International Society for Horticultural Science 4th International Symposium on Irrigation of Horticultural Crops, 1–5 September 2003, Davis, CA, USA [as yet unpublished], and Kang et al., 2003Go).

{gamma}-ray attenuation:
A related approach to the study of changes in stem water content was the use of {gamma}-ray attenuation (Brough et al., 1986Go). Although this was shown to be very sensitive, safety considerations and cost have largely limited the further application of this approach.

Sap flow
The development of reliable heat pulse and energy balance thermal sensors for sap-flow measurement in the stems of plants (Granier, 1987Go; Cohen et al., 1981Go; Cermak and Kucera, 1981Go) has opened up an alternative approach to irrigation scheduling based on measurements of sap-flow rates. Because sap-flow rates are expected to be sensitive to water deficits and especially to stomatal closure, many workers have tested the use of sap-flow measurement for irrigation scheduling and control in a diverse range of crops, including grapevine (Eastham and Gray, 1998Go; Ginestar et al., 1998aGo, bGo), fruit and olive trees (Ameglio et al., 1998Go; Fernandez et al., 2001Go; Giorio and Giorio, 2003Go; Remorini and Massai, 2003Go) and even greenhouse crops (Ehret et al., 2001Go).

Although the changes in transpiration rate that sap flow indicates are largely determined by changes in stomatal aperture, transpiration is also influenced by other environmental conditions such as humidity. Therefore changes in sap flow can occur without changes in stomatal opening. Even though rates of sap flow may vary markedly between trees as a result of differences in tree size and exposure, the general patterns of change in response to both environmental conditions and to water status are similar (Eastham and Gray, 1998Go). Appropriate sap-flow rates to use as ‘control thresholds’ may be derived by means of regular calibration measurements, especially for larger trees. Alternatively, it is at least feasible in principle to derive an irrigation scheduling algorithm that is based on an analysis of the diurnal patterns of sap flow, with midday reductions being indicative of developing water deficits (though of course diurnal fluctuations in environmental conditions can mimic such changes). Another potential problem with sap flow for precision control is that it tends to lag behind changes in transpiration rate owing to the hydraulic capacitance of the stem and other plant tissues (Wronski et al., 1985Go).

It follows that, although sap-flow measurement is well adapted for automated recording and hence potentially automated control of irrigation systems, it can be a little difficult to determine the correct control points for any crop.

Xylem cavitation
It is generally accepted (Steudle, 2001Go) that water in the xylem vessels of transpiring plants is under tension; as water deficits increase, this tension is thought to increase to such an extent that the water columns can fracture, or ‘cavitate’. Such cavitation events lead to the explosive formation of a bubble, initially containing water vapour. These cavitation events can be detected acoustically in the audio- (Milburn, 1979Go) or ultrasonic-frequencies (Tyree and Dixon, 1983Go), and the resulting embolisms may restrict water flow through the stem. Substantial evidence, though largely circumstantial, now indicates that the ultrasonic acoustic emissions (AEs) detected as plants become stressed do indeed indicate cavitation events and that AE rates can be used as an indicator of plant ‘stress’ (Tyree and Sperry, 1989Go). Nevertheless, there remain many uncertainties as it seems that at least a proportion of the AEs detected as woody tissues dry out may not be related to xylem embolisms. For example, the large numbers observed by Sandford and Grace (1985)Go as coniferous stems dried out were substantially in excess of the number of conducting tracheids present, thus suggesting a major contribution to observed AEs by the non-conducting fibres (Jones and Peña, 1986Go). Although the measurement of AEs has proved to be a powerful tool for the study of hydraulic architecture in plants, there has been little progress in adapting this measure as an indicator for irrigation scheduling. Note, however, the recent report by Yang et al. (2003)Go, who implemented a control algorithm based on the association of AE rate with transpiration rate for the precision irrigation of tomato. It is likely that the main reasons for the lack of uptake include the fact that the relationship between the number of AEs and water status changes with successive cycles of stress, and the fact that cavitation events are mostly observed during the drying phase, not during rewetting, and so cannot provide an indicator of when irrigation has been sufficient to replenish the soil water supply.

Stomatal conductance and thermal sensing
As outlined above, it appears that changes in stomatal conductance are particularly sensitive to developing water deficits in many plants and therefore potentially provide a good indicator of irrigation need in many species. It is in this area that most effort has been concentrated on the development of practical, plant-based irrigation scheduling approaches. Although stomatal conductance can be measured accurately using widely available diffusion porometers, measurements are labour-intensive and unsuitable for automation. The recognition that leaf temperature tends to increase as plants are droughted and stomata close (Raschke, 1960Go) led to a major effort in the 1970s and 1980s to develop thermal sensing methods, based on the newly developed infrared thermometers, for the detection of plant stress (see reviews by Jackson, 1982Go; Jones and Leinonen, 2003Go; Jones, 2004Go).

An early method of accounting for the rapid short-term variation in leaf temperature as radiation and wind speed vary in the field was to refer leaf temperatures to air temperature and to integrate these differences (e.g. the Stress Degree Day measure of Jackson et al., 1977Go); significant elevation of canopy temperature above air temperature was indicative of stomatal closure and water deficit stress. The method was transformed into a more practical approach following the introduction of the crop water stress index (CWSI) by Idso and colleagues (Idso et al., 1981Go; Jackson et al., 1981Go), where CWSI was obtained from the canopy temperature (Tcanopy) according to

(1)
where Tnws is a so-called non-water-stressed baseline temperature for the crop in question at the same atmospheric vapour pressure deficit, and Tdry is an independently derived temperature of a non-transpiring reference crop (Fig. 2). In this approach all temperatures are expressed as differences from air temperature so that standard relationships for Tdry and Tnws can be used. Although this approach was found to be useful in the clear arid climate of Arizona where the method was developed, it has proved to be less useful in more humid or cloudy climates where the signal-to-noise ratio is somewhat smaller (see Fig. 2; Hipps et al., 1985Go; Jones, 1999Go). In spite of its deficiencies, there has been widespread use of infrared thermometry as a tool in irrigation scheduling in many, especially arid, situations (Jackson, 1982Go; Stockle and Dugas, 1992Go; Martin et al., 1994Go), especially with the development of ‘trapezoidal’ methods involving the combination of temperature data with a visible/near infrared vegetation index (Moran et al., 1994Go).  
In order to improve the precision of the approach in more humid or low-radiation environments, Jones (1999)Go introduced the approach of using physical dry and wet reference surfaces to replace the notional Tdry and Tnws required for equation 1. A number of recent papers have shown that this approach can give reliable and sensitive indications of stomatal closure (Diaz-Espejo and Verhoef, 2002Go; Jones et al., 2002Go; Leinonen and Jones, 2004Go) and hence has the potential to be used for irrigation scheduling. The most important recent advances in the application of thermal sensing for plant ‘stress’ detection and irrigation scheduling, however, have been provided by the introduction of thermal imagery (Jones, 1999Go, 2004Go; Jones et al., 2002Go), although their expense has meant that such systems have yet to be widely used.

In addition to the use of the absolute temperature rise as stomata close, it has also been proposed that use may be made of the fact that the variance of leaf temperature increases as stomata close (Fuchs, 1990Go). Indeed, this may be a more sensitive indicator of stomatal closure than is the temperature rise (Jones, 2004Go). Again, the introduction of thermal cameras now makes the wider use of such approaches feasible, especially when combined with automated image analysis.



Automation

 

 
The most widespread use of automated irrigation scheduling systems is in the intensive horticultural, and especially the protected cropping, sector. In general, the automated systems in common use are based on simple automated timer operation, or in some cases the signal is provided by soil moisture sensors. For timer-based operation many systems simply aim to provide excess water to runoff at intervals (e.g. flood-beds or capillary matting systems), although some at least attempt to limit water application by only applying enough to replenish evaporative losses (often calculated from measured pan evaporation; Allen et al., 1999Go). Much greater sophistication is required if an objective is to improve the overall irrigation water use efficiency or to apply an RDI system. Most of the remaining automated systems currently in operation base control on soil moisture sensing; at least this approach has the potential for greater precision and improved water use efficiency.

Applications of automated plant-based sensing are largely in the developmental stage, partly because it is usually necessary to supplement the plant-stress sensing by additional information (such as evaporative demand). In principle, with high-frequency on-demand irrigation systems one could envisage a real-time control system where water supply is directly controlled by a feedback controller operated by the stress sensor itself, so that no information on the required irrigation amount is needed. For such an approach care will be necessary to take account of any lags in the plant physiological response used for the control signal.

The use of expert systems (Plant et al., 1992Go), which 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 calculation to derive a robust irrigation schedule.

Among the various plant-based sensors that have been incorporated into irrigation control systems are stem diameter gauges (Huguet et al., 1992Go), sap-flow sensors (Schmidt and Exarchou, 2000Go) and acoustic emission sensors (Yang et al., 2003Go), though there has been most interest in the application of thermal sensors. For example, Kacira and colleagues (Kacira and Ling, 2001Go; Kacira et al., 2002Go) have developed and tested on a small scale an automated irrigation controller based on thermal sensing of plant stress. Similar approaches have been applied in the field: for example, Evans et al. (2001)Go and Sadler et al. (2002)Go mounted an array of 26 infrared thermometers (IRTs) on a centre pivot irrigation system which they used to monitor irrigation efficiency, but had not developed the system to a stage where it could be used for fully automated control. Colaizzi et al. (2003)Go have tested another system that includes thermal sensing of canopy temperature on a large linear move irrigator (where the irrigator moves across the field). In another approach to the use of canopy temperature that makes use of the ‘thermal kinetic window’, Upchurch et al. (1990)Go and Mahan et al. (2000)Go have developed what they call a ‘biologically identified optimal temperature interactive console’ for the control of trickle and other irrigation systems based on canopy temperature measurements. In this direct control system, irrigation is applied as canopy temperature exceeds a crop-specific optimum. The development of thermal infrared imaging methods of irrigation control will be aided by the recent development of automated image analysis systems for extraction of the temperatures of leaf surfaces from thermal images, including shaded and sunlit leaves, soil, and other surfaces (Leinonen and Jones, 2004Go).


Conclusions

Conclusions 

 
This review has briefly considered the current state of the art and potential opportunities for use of plant-based stress sensing as the basis for irrigation scheduling and control. The advantages and disadvantages of each of these approaches are summarized in Table 1. Although plant-based sensing has several potential advantages, including a greater relevance to plant functioning than soil-based measures, these have been offset by a number of practical difficulties of implementation that have thus far limited the development of commercially successful systems. However, pressures for enhanced water use efficiency and for greater precision in irrigation systems are likely to provide a real impetus for the development of new precision irrigation scheduling systems that take account of the irrigation need of individual plants, and may well involve greater use of plant-based sensing systems.

Acknowledgements

 
The author is grateful to sponsors of various aspects of the work presented, who include the European Commission (projects: WATERUSE, contract EVKI-2000–22061, and STRESSIMAGING, Contract HPRN-CT-2002–00254).



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Figures

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Fig. 1. Generalized sensitivities of plant processes to water deficits (modified with permission from Hsiao, 1973Go).

Figure 1

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Fig. 2. (A) Illustration of the calculation of Idso's Crop Water Stress Index: CWSI=(Tcanopy–Tnws)/(Tdry–Tnws), 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.

Figure 2

 

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Source: Journal of Experimental Botany 2004 55(407):2427-2436


Table

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Table 1. A summary of the main classes of irrigation scheduling approaches, indicating their main advantages and disadvantages





Advantages


Disadvantages

I. Soil water measurement    
(a) Soil water potential (tensiometers, psychrometers, etc.) Easy to apply in practice; can be quite precise; at least water content measures indicate ‘how much’ water to apply; many commercial systems available; some sensors (especially capacitance and time domain sensors) readily automated Soil heterogeneity requires many sensors (often expensive) or extensive monitoring programme (e.g. neutron probe); selecting position that is representative of the root-zone is difficult; sensors do not generally measure water status 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’ water to apply Not as accurate as direct measurement; need 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
III. Plant ‘stress’ sensing    
(Includes both water status measurement and plant response measurement) Measures the plant stress response directly; integrates environmental effects; potentially very sensitive In general, does not indicate ‘how much’ water to apply; calibration required to determine ‘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 the most appropriate measure for many physiological processes (e.g. photosynthesis), but this argument is generally erroneous (as it ignores root–shoot signalling) All measures are subject to homeostatic regulation (especially leaf water status), therefore not sensitive (isohydric plants); sensitive to environmental conditions which can lead to 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 ({psi}) Widely accepted reference technique; most useful if estimating stem water potential (SWP), using either bagged leaves or suckers Slow and labour intensive (therefore expensive, especially for predawn measurements); unsuitable for automation
    (iii) Psychrometer ({psi}) Valuable, thermodynamically based measure of water status; can be automated Requires sophisticated equipment and high level of technical skill, yet still unreliable in the long term
    (iv) Tissue water content (RWC, leaf thickness [{gamma}- or ß-ray thickness sensors], fruit or stem diameter) Changes in tissue water content are easier to measure and automate than water potential measurements; RWC more directly related to physiological function than is total water potential in many cases; commercial micromorphometric sensors available Instrumentation generally complex or expensive, so difficult to get adequate replication; water content measures (and diameter changes) subject to same problems as other water status measures; leaf thickness sensitivity limited by lateral shrinkage
    (v) Pressure probe Can measure the pressure component of water potential which is the driving force for xylem flow and much cell function (e.g. growth) Only suitable for experimental or laboratory systems
    (vi) Xylem cavitation Can be sensitive to increasing water stress Cavitation frequency depends on stress prehistory; cavitation–water status curve shows hysteresis, with most cavitations occurring during drying, so cannot indicate successful rehydration
(b) Physiological responses Potentially more sensitive than measures of tissue (especially leaf) water status Often require sophisticated or complex equipment; require calibration to determine ‘control thresholds’
    (i) Stomatal conductance Generally a very sensitive response, except in some anisohydric species Large leaf-to-leaf variation requires much 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 large areas of crop (especially with imaging); imaging effectively averages many leaves; simple thermometers cheap and portable; well suited for monitoring purposes Canopy temperature is affected by environmental conditions as well as by stomatal aperture, so needs calibration (e.g. using wet and dry reference surfaces)
        – 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 deficit stress

Instrumentation delicate and generally expensive

Comments that relate to all methods in a section are not repeated in subsections.

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Source: Journal of Experimental Botany 2004 55(407):2427-2436


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