Journal of Plant Physiology 171 (2014) 799806

Contents lists available at ScienceDirect

Journal of Plant Physiology

journal homepage: www.elsevier.com/locate/jplph

Electrophysiological assessment of water stress in fruit-bearing

woody plants
Liliana Ros-Rojas a , Franco Tapia b , Luis A. Gurovich a,
a
b

Facultad de Agronoma e Ingeniera Forestal, Ponticia Universidad Catlica de Chile, Chile

Universidad de Santiago de Chile, Chile

a r t i c l e

i n f o

Article history:
Received 6 December 2013
Received in revised form 16 February 2014
Accepted 20 February 2014
Available online 4 March 2014
Keywords:
Persea americana
Plant electric signal transmission
Plant electrophysiology
Prunus domestica
Water stress

a b s t r a c t
Development and evaluation of a real-time plant water stress sensor, based on the electrophysiological
behavior of fruit-bearing woody plants is presented. Continuous electric potentials are measured in tree
trunks for different irrigation schedules, inducing variable water stress conditions; results are discussed
in relation to soil water content and micro-atmospheric evaporative demand, determined continuously
by conventional sensors, correlating this information with tree electric potential measurements.
Systematic and differentiable patterns of electric potentials for water-stressed and no-stressed trees
in 2 fruit species are presented. Early detection and recovery dynamics of water stress conditions can
also be monitored with these electrophysiology sensors, which enable continuous and non-destructive
measurements for efcient irrigation scheduling throughout the year.
The experiment is developed under controlled conditions, in Faraday cages located at a greenhouse
area, both in Persea americana and Prunus domestica plants. Soil moisture evolution is controlled using
capacitance sensors and solar radiation, temperature, relative humidity, wind intensity and direction are
continuously registered with accurate weather sensors, in a micro-agrometeorological automatic station
located at the experimental site.
The electrophysiological sensor has two stainless steel electrodes (measuring/reference), inserted on
the stem; a high precision Keithley 2701 digital multimeter is used to measure plant electrical signals; an
algorithm written in MatLab , allows correlating the signal to environmental variables. An electric cyclic
behavior is observed (circadian cycle) in the experimental plants. For non-irrigated plants, the electrical
signal shows a time positive slope and then, a negative slope after restarting irrigation throughout a rather
extended recovery process, before reaching a stable electrical signal with zero slope. Well-watered plants
presented a continuous signal with daily maximum and a minimum EP of similar magnitude in time, with
zero slope. This plant electrical behavior is proposed for the development of a sensor measuring real-time
plant water status.
2014 Elsevier GmbH. Open access under CC BY-NC-ND license.

Introduction
Plant bioelectric activity in response to different stimuli was
originally reported by Burdon-Sanderson (1872) and Darwin
(1896). Plants have developed several paths for electric signal transmission between individual cells, tissues and organs, in order to
adapt and optimize its physiological processes to sudden changes
in environmental stress conditions (Fromm, 2006; Mancuso and

Abbreviations: AP, action potential (mV); EP, electric potential (mV); SD,
standard deviation; VP, variation potential (mV); , volumetric soil water content
(%).
Corresponding author at: P.O. Box 30622, Santiago, Chile. Fax: +56 226864130.
E-mail address: lgurovic@puc.cl (L.A. Gurovich).
http://dx.doi.org/10.1016/j.jplph.2014.02.005
0176-1617/ 2014 Elsevier GmbH. Open access under CC BY-NC-ND license.

Mugnai, 2006; Fromm and Lautner, 2007; Wang et al., 2009; Yan
et al., 2009; Sukhov et al., 2011; Gurovich, 2012; Volkov, 2012).
Sibaoka (1966), working with Mimosa pudica, reported the
onset of electric activity, equivalent to an action potential (AP),
and Pickard (1973), experimenting with higher plants, described
similar results. Different studies have been published on electric
signaling in plant cells and tissues, resulting from biotic and abiotic
stimuli, resembling an electrical replication of the specic stimulation (Fromm, 2006). AP is generated in plants in response to a
stimulus larger than a certain threshold, in the form of an all or nothing response, as documented by several authors (Datta and Palit,
2004; Lautner et al., 2005; Gil et al., 2008; Volkov et al., 2012; Wang
et al., 2009; Oyarce and Gurovich, 2010). Also, variation potentials
(VPs) have been reported in plants, characterized as an electric
response to stimuli smaller that the specic AP threshold, being

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its magnitude a function of its intensity (Yan et al., 2009). It has

been suggested that VP transmission rates are related to xylematic
turgor conditions and VP is also known as short wave potential
(Stahlberg and Cosgrove, 1995; Gensler and Yan, 1998; Stahlberg
et al., 2005).
Electrical signal generation and transmission in plants has been
documented in relation to modications in light intensity (Volkov
and Haack, 1995; Trebacz et al., 2006; Cabral et al., 2011; Volkov
et al., 2012), osmotic pressure (Schroeder and Hedrich, 1989), temperature (Volkov et al., 2007; Wang et al., 2009; Cabral et al.,
2011; Kai et al., 2011), mechanical wounds and insect activity
(Brenner et al., 2006), changes in cytoplasmic Ca2+ concentrations (Knight et al., 1991; Fromm and Spanswick, 1993; Shepherd,
2005), changes in respiration, photosynthesis and phototropism
(blue light) (Lautner et al., 2005; Pavlovic and Mancuso, 2011),
ower induction (Parimalan et al., 2005; Wagner et al., 2006,2012),
defense mechanisms to herbivore damages (Volkov and Ranatunga,
2006; Fromm and Lautner, 2007; Heil and Ton, 2008), mechanical stimulation (Braam, 2005; Volkov and Ranatunga, 2006) and
soil water availability (Fromm and Fei, 1998; Brenner et al., 2006;
Gurovich and Hermosilla, 2009; Oyarce and Gurovich, 2010).
Several reviews related to plant electric responses to different
stimuli have been published by Fromm and Lautner (2007), Baluska
and Mancuso (2009) and Gurovich (2012). Electric signals can be
transmitted between plant organs and tissues because changes
in the trans - membrane potential induced by a specic stimulus
create a depolarization wave, propagating through plasmatic membranes of excitable adjacent cells (Volkov and Ranatunga, 2006),
enabling plants to generate and coordinate physiological activity
at signicant distances (Lautner et al., 2005). Transmission rates
up to 40 m s1 for AP have been reported by Volkov and Brown
(2004), Oyarce and Gurovich (2010). For VP transmission, rate
ranges between 0.1 and 10 mm s1 have been measured (Stahlberg
and Cosgrove, 1995, 1997; Koziolek et al., 2003; Stahlberg et al.,
2005).
Electric signal transmission in plants takes place mainly through
cellular connections in the symplastic phloematic continuum (Bose,
1926; Cosgrove and Hedrich, 1991; Fromm and Fei, 1998; Lautner
et al., 2005; Van Bel and Ehlers, 2005). Essentially, it is an electrochemically activated communication system, operating as a simple
neuronal net (Mancuso, 1999; Volkov et al., 2007; Zimmermann
et al., 2009).
Evidences of electric signal transmission between distant plant
organs have been documented by Fromm and Fei (1998), reporting electric potential (EP) rates of 1 cm s1 , which induced stomata
closing before a reduction of leaf turgor could be measured in
Zea mays plants under water stress. Also, Grams et al. (2007)
reported EP differences of 50 mV and electric signal transmission
rates of 1 cm s1 , associated to an increment in stomata conductance and CO2 absorption, which starts 60 s after rehydration. These
results indicate that signals different to hydraulic (Mullendore et al.,
2010) or hormone signaling (Rodrguez-Gamir et al., 2011), must
be present as an information pathway between plant organs, to
account for these fast physiological responses to changes in water
availability, dened as a dynamic steady state of the soil water ow
to roots and the evapo-transpirative leaf water ow to the atmosphere (Koziolek et al., 2003; Fromm, 2006; Gibert et al., 2003).
Most studies on EP in plants have been carried on under
laboratory or greenhouse controlled conditions, mainly in succulent plants (Mwesigwa et al., 2000; Volkov et al., 2007, 2009,
2012). In woody species, like Vitis vinfera, Mancuso (1999),
measured signicant EP changes as a result of leaf and shoot
aming injury and mechanical wounds, reporting also a detection of the resulting electric signal in distant plant organs, a
short time after the injury event. Gibert et al. (2006) reported
a signicant correlation between EP and sap ow in Populus

trichocarpa trees, concluding that the electric signals measured

are propagated through the phloem, at short or long distances,
modifying photosynthetic rates. Systematic electric signaling in
fruit-bearing woody species, induced by alternate light/dark cycles
and soil water decits, have been reported by Gil et al. (2008,
2009), Gurovich and Hermosilla (2009), Oyarce and Gurovich
(2011).
This research presents evidence on systematic EP variations in
Prunus domestica and Persea americana trees for alternate cycles
of full irrigation and water restriction periods, in association
with alternate micro environmental conditions (soil water content
depletion and evapotranspiration rates). Electrophysiological sensors developed for this research enable real-time EP measurements,
to be used as a non-destructive and continuously agronomic tool for
irrigation scheduling in fruit-bearing woody species, as an alternative to discrete xylematic pressure measurements or other soil and
plant stress assessment techniques available, which are time consuming and highly dependent on the operators skill, to accurately
represent the actual plant water status.
Materials and methods
Experimental setup and plant material
The experiment was located at the greenhouse area adjacent
to the School of Agriculture and Forestry, Ponticia Universidad
Catolica de Chile in Santiago, Chile. Two independent Faraday cages
are isolated from environmental electromagnetic elds, using a
tested grounded connection. In each cage, ten 3-years-old avocado plants (Persea americana Mill., cv. Hass, grafted on Mexicola
rootstock) and plum (Prunus domestica L., cv. DAgen, grafted on
Mariana 2624 rootstock), are grown in individual 20 L containers
in easily drainable 50% peat moss/50% perlite (Prunus domestica L.)
and 100% perlite (Persea americana Mill.) substrates respectively,
enabling to create water decit conditions in short periods of time.
An independent, programmable irrigation system is available on
each Faraday cage, to create differentially controlled water supply
treatments, in order to evaluate plant EP responses to alternated
water availability conditions.
Electric connections in plants
Two electrodes are inserted on each lignied trunk at 20 cm (reference electrode) and 30 cm (measuring electrode) above the graft
(Fig. 1), according to our previous experimental results (Gurovich
and Hermosilla, 2009). Each electrode consists of a solid stainless
steel 304 SS type 316 rod, 3.18 cm long and 0.2 cm diameter, connected to a 0.5 mm cooper conductor, shielded from air relative
humidity uctuations by a exible self-adhesive plastic ribbon.
Each electric circuit is connected to an ampliermulti voltmeter (Keithley, model 2701), provided with two 20 channel
multiplexer plates model 7700 (Fig. 2), with the positive terminal for the measurement electrode and the negative terminal
for the reference electrode, on its high (H) y low (L) insertion
points, respectively. Registration of EP measurements, using the
EXCELinx utility software, is made every 10 s and each measurement lasts 0.1 ms. EP information registered is analyzed in graphic
format using a MatLab algorithm, developed for this research.
Simultaneously to EP measurements, electric capacitance sensors (Decagon model 10HS), located within each plant container
peat moss/perlite or perlite substrate, enables real-time volumetric
soil water measurements (, %); also, micro-environmental parameters are registered using sensors integrated into an automatic
meteorological station (radiometer, hygrometer, thermometer,
wind speed and direction sensors), located between the two

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801

Fig. 1. Electrode insertions in the tree trunk.

Faraday cages; data is transmitted through the Internet to the

WiseConn S. A. server, and analyzed using the DropControl
software, enabling a graphic display of each parameter and the calculation of hourly ETpotential values, by using an internal algorithm
based on Penman-Monteith model, modied by FAO (Allen et al.,
1998). Plant EP (the difference between EP values simultaneously
measured at both electrodes) are correlated to  and to meteorological parameters (solar radiation sr, W m2 , air temperature T, C)
to measure plant electric behavior in response to variations in the
dynamic steady state water availability conditions.
A water restriction experiment is reported, lasting from March
15 to April 11, 2013, period characterized by a high atmospheric

evaporative demand (4.41 mm d1 ; standard deviation (sd): 0.79).

A ve plant subset, on each Faraday cage, is irrigated every 12 h
adding 2.4 L tap water to each container, throughout the experiment, keeping  at 43.1% (0.5%) for Prunus domestica and  at 30.1%
(3.8%) for Persea americana, respectively. A ve plant subset in
each species is kept without irrigation between March 28 to April 3,
when visual water stress symptoms are evident in the early morning hours and  reaches 16.3% and 8.3% for Prunus domestica and
Persea americana, respectively. Irrigation every 12 h is again supplied to all experimental plants to start a recovery period, lasting
until April 11.

Results

Fig. 2. Keithley 2701 amplier multi voltmeter connections.

EP measurements and environmental conditions in Prunus

domestica plants are recorded from March 15 to April 11, 2013
(Fig. 3a (variations of solar radiation, in the range 0660 W m2 );
Fig. 3b (air temperature, in a range 632 C); Fig. 3c and d (substrate
volumetric water content ). EP values for each plant subset (irrigated twice a day and non-irrigated between March 28 and April 3,
respectively) are measured in a representative plant. EP values presented in Fig. 3c and d indicates the existence of a differential plant
electrical response as related to solar radiation and air temperature
variations; likewise, the maximum daily EP values, for plants subject to water restriction between March 28 and April 03 (blue band
in Fig. 3c and d) are signicantly higher than the maximum daily
EP values measured in non-restricted water plants.
As the substrate dries, the maximum daily EP value increases
from 18.8 mV to 92.9 mV, detected at 11:29 am (sd 0.26). Minimum EP values, detected at 6:45 pm, (sd 1.08), range 7.09 mV at
the onset of the non-irrigated period and 57.13 mV, just before
the rst irrigation is supplied, ending the water restriction period,
on April 3 at 00:30. The maximum and minimum daily EP values
increase 1.4 mV  1 for the period of water restriction, in the range
47.8% >  > 16.3%. When plants are kept in a twice daily irrigation
regime, with constant , the temporary variation in the maximum
and minimum EP daily values vary between 18.44 mV, (sd 5.22)
and 48.45 mV (sd 4.66), events taking place at 9:48 am (sd 2.05)
and 6:43 pm (sd 1.43), respectively.

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Fig. 3. PE evolution of Prunus domestica for two conditions of soil humidity. (a) Radiation (W m2 ). (b) Temperature ( C). (c) Unirrigated plants during 5 consecutive days.
(d) Plants with 2 daily irrigations.

Plant electrical response presents a periodical pattern, with a

daily average oscillation of 20 mV, which increases to 80 mV, for
plants without irrigation. According to the information presented
in Fig. 3, plant electrical response is also affected by the variation
on environmental factors, like air temperature and solar radiation,
which decreased during the period of water restriction at a rate of

3.3 C day1 and 140 W m2 day1 , respectively (Fig. 4a). Differential electrical responses for watered plants (gray line) and water
restricted plants (black line) indicate the specic effect of water
availability on plant EP (Fig. 4b), when the signal is analyzed using
the wavelet decomposition method (Lewalle et al., 2007) enabling to
isolate a rst-order-numeric decomposition of the electrical signal,

Fig. 4. (a) Air temperature and solar radiation rates. (b) First-order-decomposition of EP variations for Prunus domestica in two conditions of soil water content: unirrigated
plants during 5 days (black line), and irrigated plants every 12 h (gray line).

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803

Fig. 5. (a) EP frequency associated to the circadian cycle for Prunus domestica, after removing the rst-order-decomposition of the signal. (b) Spectral analysis for the period
evaluated, frequency 1 Hz, equivalent to one day.

Fig. 6. PE Evolution of Persea americana for two conditions of soil humidity. (a) Radiation (W m2 ). (b) Temperature ( C). (c) Unirrigated plants during 5 consecutive days.
(d) Irrigated plants every 12 h. Same period and treatment applied to Prunus domestica (see Fig. 3).

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in order to remove the circadian cycle effect (McClung, 2001) and to

quantify other environmental effects. EP response of Prunus plants
under specic environmental conditions of radiation, temperature
and water availability, suggests possible systemic plant coordination, by means of electrical signaling taking place between different
plant organs, during the development of plant water stress, possibly determining the magnitude and intensity of its physiological
responses.
Fig. 5a shows EP oscillatory behavior during the whole
experimental period, showing a clear circadian frequency (high
frequency), using Sadowsky (1996) wavelet method to analyze EP
values for a representative experimental Prunus plant (Fig. 5b) in
four time intervals considered: irrigated plants, March 1528 and
April 0511 (green band), plants with water restriction, March
28April 03 (gray band) and plants with water restriction but in
a recovery stage, April 0305 of 2013 (red band); the circadian
behavior occurs in irrigated and non-irrigated plants in the four
periods.
From April 3, when irrigation water is re-supplied to plants previously subjected to water restriction, a recovery period begins (red
band), with an irregular electrical response to sudden rehydration;
this irregular electrical behavior is not observed in plants under no
water restriction. Two days after irrigation is resumed, and the soil
is kept continuously at  = 43.31, plant electrical behavior of both
plant sets is almost totally stabilized, probably indicating the end
of plant water stress condition.
In Fig. 5b a power spectrum analysis of EP (Percival and Walden,
1993) shows a specic peak in frequency domain. This fundamental harmonic frequency is approximately 1 Hz, equivalent to a
one-day period; the subsequent peaks correspond to an harmonic
oscillation, associated to a signal distortion of the fundamental
pattern, allowing us to infer that the stress-recovery mechanism
during the circadian cycle varies as a function of water stress intensity, as the plant recovers from the water restriction previously
imposed.
Similarly to Prunus, Persea plants respond electrically to conditions of water restriction, with a systematic increment on its
maximum and minimum daily EP values, as the substrate dries
out (Fig. 6). Maximum daily EP values increase from 14.56 mV to
52.5 mV, from the onset to the end of the water restriction period,
respectively (Fig. 6c); these EP values are detected at 6:50 pm (sd
1.54). EP minimum daily values, detected at 11:31 am (sd 2.16),
range from 2.62 mV at the beginning of the non-irrigated period to
41.4 mV, just before the April 3 at 12:30 pm irrigation event. Maximum and minimum EP values increase 4.02 mV  1 under water
restriction conditions.

Fig. 7. First-order-decomposition of the PE variations for Persea americana for two

conditions of soil water content: unirrigated plants during 5 days (black line), and
irrigated plants every 12 h (gray line).

A cyclic variation in EP maximum and minimum values is

observed when the plants are maintained with constant  (Fig. 6d);
the cycle is determined only by the variations of solar radiation
and temperature (Fig. 6a). However, there is a clear inuence of
water restriction on Persea americana EP behavior, when irrigated
and not irrigated plants EP responses are compared (Fig. 7). The
periodical EP oscillation, corresponding to the twice daily watered
plants (Fig. 7, gray line), is kept under 20 mV, while a progressive
EP increment, exceeding 40 mV, is measured in non-watered plants
(black line).
EP varies throughout the day, both under a twice daily irrigation
schedule and also, when water supply is interrupted during several
consecutive days. This EP behavior has been related to light and
darkness cycles, as reported by Gurovich and Hermosilla (2009);
plants present also an EP circadian cycle (Oyarce and Gurovich,
2010), showing regular EP value amplitudes between consecutive
days for both species (Fig. 8), even under no water restriction.
Maximum daily EP for Prunus (red line in Fig. 8a) typically occurs
at 11:05 am (sd 0.07), while for Persea (black line in Fig. 8a) this
maximum daily PE is measured by late afternoon, at 7:21 pm (sd
0.22). This difference (, Fig. 8b) between both species, shows a
79 h magnitude.
EP behavior throughout the day is different in both species
(Fig. 8). For Prunus, irrigation at mid-day does not affect the daily
PE cycle; the maximum PE occurs before the irrigation event; for
Persea, mid-day irrigation determines a change in the sequence
of PE values, with a decrease occurring immediately after the
irrigation event, and a further increase, until a maximum daily PE
is reached by late afternoon. During the water restriction period,
when the substrate dries out, EP behavior in both species is altered;
the daily amplitude (difference between the maximum and minimum EP) in Prunus (Fig. 3c), decreases from 13.21 mV to 8.48 mV, at

Fig. 8. (a) Mismatch of the circadian cycles, represented by the electrical activity of the Prunus domestica (red line) and Persea americana (black line) plants under constant
irrigation (every 12 h). (b) Daily mismatch () maximum PE through time. (For interpretation of the references to colour in this gure legend, the reader is referred to the
web version of this article.)

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805

Fig. 9. Differences in EP circadian cycles. (a) Prunus domestica plants and (b) Persea americana plants, under two water availability conditions. In both gures, red line: drought
treatment and blue line: no drought. (For interpretation of the references to colour in this gure legend, the reader is referred to the web version of this article.)

the beginning of the non-irrigated period and before the irrigation

restitution, respectively.
In the case of Persea (Fig. 6c) EP amplitude variations throughout the day are not conclusive, as the increased EP differences are
only present in the last 2 days of the period, when plants are kept
continuously without water supply.
On April 3, just before irrigation is restored after the water
restriction period, maximum and minimum PE values for Prunus
occurs at the same time, both for plants with and without water
restriction (Fig. 9a); for Persea, there is a signicant difference in the
maximum and minimum EP timing for irrigated and non-irrigated
plants (Fig. 9b). These inter-specic differences in the electrical
behavior occur every day throughout the experiment; thus, to adequately interpret EP measurements as indicators of early water
stress, it is necessary to quantitatively consider these differences,
correlating EP species behavior to other physiological indicators,
like stomata conductance (Volkov, 2012).
Discussion
Both Prunus and Persea plants respond electrically to conditions
of water restriction, with increased maximum and minimum EP
daily values, throughout the period when water supply id restricted
and the substrate progressively dries out due to evapotranspiration.
Differential EP behavior, due to solar radiation and temperature
variations, are also detected between this two species, suggesting
a possible coordination, by means of electrical signals, among its
different organs (Volkov et al., 2007; Zimmermann et al., 2009).
Plant adaptation to different water availability conditions, possibly
is determining the magnitude and intensity of their physiological
responses (Gurovich and Hermosilla, 2009; Oyarce and Gurovich,
2010).
Our results indicate some differential EP behavior, both when
plants are irrigated twice daily, and also when water supply is
interrupted during several consecutive days; similar effect of alternate sunlight and darkness conditions upon EP behavior has been
reported by Gurovich and Hermosilla (2009), Oyarce and Gurovich
(2010), which also reported a circadian cycle.
A low frequency analysis of EP data, for the rst-orderdecomposition of the signal, shows a signicantly higher electrical
signal gradient for water restricted plants, as compared to irrigated
plants; this differential behavior could be used as an early indicator
of the water stress. However, this type of analysis must include data
for time intervals larger than a single day, in order to determine if
the variation of EP responses corresponds to an actual water stress,
instead of to a different environmental factor.

EP differential behavior reported in this work, both for Prunus

and Persea, indicate that water stress in woody plants can be characterized by continuous increments of EP. Results from this study
are similar to those published by Fromm and Fei (1998), Brenner
et al. (2006), Gurovich and Hermosilla (2009), Oyarce and Gurovich
(2010), providing experimental support to the hypothesis that realtime EP measurements can be used as early indicators of plant
water status. However, intrinsic differences in phenotypic (foliar
index area) and physiological (stomata conductance) characteristics of plant species determine the absolute magnitude of EP
increments, under the same atmospheric evaporative demand (Gil
et al., 2009; Egea et al., 2010). Therefore, specic models of electrical response are necessary for a precise interpretation of EP values
as valid indicators of water stress in fruit-bearing trees.

Conclusions
Results presented in this work indicate the possibility to use
plant electrophysiological behavior to develop an agronomy sensor for real-time measurements of plant water status, for its use
on accurate irrigation scheduling of fruit trees, if an interpretation model on EP responses to water availability can be dened
for different species.
Using a low-frequency analysis on the electrical signal, it is feasible to separate the effects of environmental stimuli from the
specic effect of water restriction in plant electrical response. Also,
the use of an high-frequency signal analysis and its corresponding harmonious, can detect a temporary correlation of EP with
the circadian cycle for each plant species, indicating that plant
EP behavior is probably determined to a large extent by its morphological and physiological characteristics, interacting with its
micro-environment.
High and low frequency separation analysis enable us to quantify the effect of the substrate drying process through temporary
variations of daily EP maximum and minimum values, by comparing similar measurements taken in non-water stressed plants.
Also, during the recovery period, when irrigation is resumed, plant
EP behavior indicates that there is an additional period of at least
3 days, on which previously non-irrigated plants still manifest a
different electrical behavior, as compared to non-stressed plants,
indicating that the process of water stress recovery can be a rather
long period, lasting several days after full irrigation is resumed.
We observed a differential EP circadian cycle behavior for both
species: while micro-environmental or water availability modications determine a time-shift on the cycle in Persea, the cycle is not

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altered in Prunus plants; physiological causes and implications of

this differential behavior will become important in future studies.
Acknowledgements
Authors acknowledge the collaboration of Fondo de Innovacin
Agraria FIA, Ministry of Agriculture, Chile (project PYT 2011-0038)
and the Wiseconn S. A. Company for its electronic and software
support.
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