Agriculture, Ecosystems and Environment 141 (2011) 271286

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Agriculture, Ecosystems and Environment

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Review

Stressed food The impact of abiotic environmental stresses on crop quality

Yunxia Wang a , Michael Frei b,
a
Key Laboratory of Crop Genetics & Physiology of Jiangsu Province, Yangzhou University, 12 Wenhui Road, 225009 Yangzhou, China
b
Institute of Crop Science and Resource Conservation (INRES) Plant Nutrition, University of Bonn, Karlrobert-Kreiten Strasse 13, D-53115 Bonn, Germany

a r t i c l e i n f o a b s t r a c t

Article history: Abiotic environmental stresses negatively impact crop productivity and are major constraints to global
Received 7 January 2011 food security. As a consequence of global change, certain stress factors such as heat, drought, salinity,
Received in revised form 25 March 2011 tropospheric ozone, and excess UV radiation might become even more prevalent in the coming decades.
Accepted 28 March 2011
While the negative impact of these stresses on crop yields is obvious, their effects on crop quality are
Available online 6 May 2011
less recognized. Exposure to environmental stress induces numerous physiological stress reactions in
plants that can alter the chemical composition of crops and thus the quality of the harvested products.
Keywords:
Literature on the impact of abiotic environmental stresses on crop quality falls into seven categories of
Global change
Crop production
quality parameters: protein, lipids, non-structural carbohydrates, minerals, antioxidants, feed value for
Nutritional value ruminant herbivores, and physical/sensory traits. Apart from summarizing net effects on these quality
Stress adaptation parameters, this review intends to elucidate physiological mechanisms leading to the observed changes in
Food security crop quality. All categories of traits are signicantly affected by abiotic environmental stresses, resulting
Crop quality in both positive and negative changes in crop quality. The overall effect of a certain stress factor is often
dependent on numerous interacting factors such as the timing of stress application, the intensity of
the stress, and the crop species. In spite of these confounding elements, this review identies some
common patterns of stress response, such as a tendency towards increasing concentrations in protein and
antioxidants in stressed crops, and a loss in quality in terms of feed value, starch and lipid concentration, or
physical/sensory traits. This information might help agronomists and crop breeders to develop strategies
to produce higher quality crops in stress environments.
2011 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
2. Protein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
3. Lipids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
4. Non-structural carbohydrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
5. Minerals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
6. Antioxidants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 276
7. Feed value . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
8. Physical and sensory traits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
9. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
10. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281

1. Introduction (e.g. Ashmore et al., 2006; Battisti and Naylor, 2009). Abiotic stress
factors such as drought, heat, soil salinity, tropospheric ozone and
Many recent studies have identied environmental stresses as excess UV radiation are already causing signicant agricultural
major threats to global food security in the twenty-rst century yield losses and will become even more prevalent in the coming
decades due to the effects of global change (Ashmore et al., 2006;
Ortiz et al., 2008; Battisti and Naylor, 2009; Feng and Kobayashi,
Corresponding author. Tel.: +49 228 732851; fax: +49 228 731695. 2009; Fuhrer, 2009; Wassmann et al., 2009a). Tropical and sub-
E-mail addresses: yxwang@yzu.edu.cn (Y. Wang), mfrei@uni-bonn.de (M. Frei). tropical developing countries in Asia and Africa that face high

0167-8809/$ see front matter 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.agee.2011.03.017
272 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286

population growth are at a particularly high risk of food shortage Table 1

Effects of environmental stress factors on the protein concentration of the harvested
caused by these stress factors (Lobell et al., 2008; Wassmann et al.,
fractions of crops.
2009a).
Agricultural yield losses due to abiotic environmental stresses Crop species Effect References
are obvious and have been well documented. The awareness of Drought
the growing impact of environmental stress has lead to worldwide Barley, Hordeum vulgare L. Savin and Nicolas (1996)
efforts in adapting agricultural production to such adverse environ- Corn, Zea mays L. Oktem (2008)
Peanut, Arachis hypogeal L. Dwivedi et al. (1996)
mental conditions (Lobell et al., 2008; Ortiz et al., 2008; Wassmann
Potatoe, Solanum tuberosum L. Teixeira and Pereira (2007)
et al., 2009b). Proposed strategies to face these challenges include Soybean, Glycine max (L.) Merr. Rotundo and Westgate
improved agronomic management and the breeding of novel crop (2009)
genotypes adapted to stress environments. For example, molecu- Wheat, Triticum aestivum L. Altenbach et al. (2003),
Gooding et al. (2003),
lar breeding projects have been initiated to adapt novel varieties of
Rharrabti et al. (2003),
major cereal crops to heat (Pinto et al., 2010), drought (Araus et al., Ozturk and Aydin (2004),
2008; Fleury et al., 2010), salinity (Ren et al., 2005), and tropo- Guttieri et al. (2005),
spheric ozone (Frei et al., 2008). The focus of most agronomic and Asseng and Milroy (2006),
breeding efforts has been on mitigating quantitative yield losses Weightman et al. (2008),
Saint Pierre et al. (2008),
caused by environmental stress factors.
and Flagella et al. (2010)
Far less attention has been devoted to the impact of abi- Heat
otic environmental stresses on crop quality. Exposure to stress Barley, Hordeum vulgare L. Savin and Nicolas (1996)
leads to numerous physiological changes in crop plants, such as and Pettersson and
Eckersten (2007)
changes in the photosynthetic gas exchange and assimilate translo-
Rapeseed, Brassica napus L. Triboi and Triboi-Blondel
cation (Martin and Ruiztorres, 1992; Morgan et al., 2004), altered (2002)
water uptake and evapotranspiration (Rivelli et al., 2002; Katerji Rice, Oryza sativa L. Lin et al. (2010)
et al., 2010), effects on nutrient uptake and translocation (Hu and Sunower, Helianthus annuus L. Triboi and Triboi-Blondel
Schmidhalter, 2005; Sanchez-Rodriguez et al., 2010), antioxidant (2002)
Soybean, Glycine max (L.) Merr. Rotundo and Westgate
reactions (Blokhina et al., 2003; Apel and Hirt, 2004), programmed
(2009)
cell death (Kangasjrvi et al., 2005), and altered gene expression Wheat, Triticum aestivum L. Stone et al. (1997),
and enzyme activity (Yamakawa et al., 2007; Guo et al., 2009; Frei Altenbach et al. (2003),
et al., 2010b). If such processes occur over extended periods of time Dupont et al. (2006), and
Asseng and Milroy (2006)
due to chronic exposure to stress, they are very likely to affect the
Salinity
chemical composition of crops, and thus the quality of agricultural Potatoe, Solanum tuberosum L. Teixeira and Pereira (2007)
products. Tropospheric ozone
This review summarizes scientic literature on the effects of ve Bahiagrass, Paspalum notatum Flugge Muntifering et al. (2000)
abiotic stress factors (drought, heat, salinity, tropospheric ozone, Common Bean, Phaseolus vulgaris L. Iriti et al. (2009)
Corn, Zea mays L. Garcia et al. (1983)
and UV radiation) on the quality of agricultural products. Stress
Mustard, Brassica campestris L. Singh et al. (2009)
is dened as a growth condition, in which one of these factors Peanut, Arachis hypogeal L. Burkey et al. (2007)
is elevated relative to a non-stress control treatment. It considers Rapeseed, Brassica napus L. Bosac et al. (1998) and
only studies reporting data on the quality of harvested food prod- Ollerenshaw et al. (1999)
Soybean, Glycine max (L.) Merr. Kress and Miller (1983),
ucts, while ignoring all those studies dealing with vegetative plant
Mulchi et al. (1988), and
materials and premature crops. Apart from summarizing the net Heagle et al. (1998)
effects on quality, we also attempt to elucidate the physiological Wheat, Triticum aestivum L. Kress and Miller (1983),
factors leading to the observed changes in crop quality. The review Mulchi et al. (1986),
is structured into seven chapters, each of which explores the effects Vandermeiren et al. (1992),
Fuhrer et al. (1989, 1990,
of environmental stresses on a specic quality parameter: protein,
1992), Finnan et al. (1996),
lipids, non-structural carbohydrates, minerals, antioxidants, feed Rudorff et al. (1996), Pleijel
value, and physical/sensory quality traits. et al. (1991, 1997, 1998,
1999, 2006), Pleijel (1998),
Feng et al. (2008), and
Piikki et al. (2008)
2. Protein
() Stress increased protein concentration; () stress decreased protein concentra-
tion; () stress had no clear effect on protein concentration.
Food crops are an important source of dietary protein for
humans and animals. Although the protein concentration of com-
monly used crops is only 1030% of the total dry weight (Triboi and tration in wheat grains and soybean seeds increased as a result of
Triboi-Blondel, 2002), it plays a signicant role in determining the all investigated stresses, including drought, heat and tropospheric
nutritional quality of many crops, particularly cereals and legumes. ozone. In the case of rapeseed, heat stress increased the protein
This review addresses only the inuence of major abiotic environ- concentration (Triboi and Triboi-Blondel, 2002), but ozone had
mental stresses on total protein (crude protein) concentration of the opposite effect (Bosac et al., 1998; Ollerenshaw et al., 1999).
harvested fractions of food crops; changes in protein composition Apart from the type of stress or the crop species, the timing of
or protein quality are not further explored here readers inter- stress occurrence was an important factor determining the effect
ested in this subject may be referred to the review by Dupont and on protein concentration of the harvested fractions, which may par-
Altenbach (2003). tially explain inconsistent effects in studies dealing with the same
Protein concentrations of crops are affected by both genetic stress and crop species (Rotundo and Westgate, 2009). For example,
and environmental factors. Environmental stresses or unfavorable Dwivedi et al. (1996) found that mid-season drought had no effect
growth conditions usually induce higher protein concentration in on the protein concentration of peanut, while end-season drought
the harvested fractions of crops, with only a few studies showing signicantly increased the total protein concentration. In the case
no effect or lower protein concentration (Table 1). Protein concen- of rice, high temperatures during grain lling increased the accu-
 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286
mulation of all classes of storage proteins at the early lling stage,
but decreased the accumulation of prolamins at maturation (Lin
et al., 2010).
Compared with other stresses, the effects of ozone pollution
were more complex and varied with ozone concentration, exposure
time, species or cultivars, and the inuences of other environmen-
tal factors such as carbon dioxide. The majority of studies found
that ozone increased the protein concentration of wheat grains
and soybean seeds, despite experimental differences in ambient
ozone or elevated ozone treatments (Table 1). Ambient ozone con-
centration increased the crude protein concentration of bahiagrass
and common bean compared to charcoal ltered air (Muntifering
et al., 2000; Iriti et al., 2009), but decreased that of mustard (Singh
et al., 2009). In contrast, twice-ambient ozone concentration had
no effect on protein concentration in peanut (Burkey et al., 2007).
In the case of rapeseed, different cultivars of the same species
responded differently to ozone stress in terms of protein concen-
tration (Bosac et al., 1998).
Rising carbon dioxide concentration is an important environ-
mental factor for the quality of food crops that is expected to
accompany rising ozone concentrations (Loladze, 2002; Taub et al.,
2008; DaMatta et al., 2009). Elevated carbon dioxide concentra-
tions tend to counteract ozone effects on yield and crop quality, and
vice versa. The nature of such interactions depends on the species
and the gas concentrations. In potato, tuber protein concentrations
were decreased by elevated carbon dioxide under both high and
low levels of ozone. However, in soybean, elevated carbon diox-
ide increased protein concentrations under high ozone, but had
the opposite effect under low ozone (Taub et al., 2008). A meta-
analysis on wheat by Feng et al. (2008) demonstrated that elevated
ozone increased the grain protein concentration but decreased the
protein yield (the absolute amount of protein produced). Similar
results were found by Piikki et al. (2008); in addition, they found
that elevated carbon dioxide had the opposite effect on grain pro-
tein compared to ozone, but the changes in grain quality due to
ozone were not simply offset by elevated carbon dioxide. In peanut,
elevated carbon dioxide mitigated negative ozone effects on yields
to varying extents depending on the concentrations of the two
gases, but neither carbon dioxide nor ozone affected the protein
concentrations of seeds (Burkey et al., 2007; Tu et al., 2009).
Although different stresses impose different physiological reac-
tions in crops, the common negative effects on dry matter
accumulation have been suggested by many researchers as a main
factor related to higher protein concentrations. In most cases, the
nitrogen harvest index, representing the proportion of nitrogen
contained in the grain relative to total above-ground nitrogen con-
tent (Cox et al., 1986), was less affected by stresses than dry matter
harvest index, which resulted in higher protein concentration in
the harvest fractions of crops (Gooding et al., 2003; Asseng and
Milroy, 2006; Weightman et al., 2008). Pleijel et al. (1999) investi-
gated the grain protein accumulation in relation to the grain yield of
spring wheat grown in open-top chambers with different concen-
trations of ozone and carbon dioxide across four different countries.
They found a negative linear correlation between grain yield and
grain protein concentration. Thus the changes in grain protein con-
centration could be explained largely by a simple concentration
effect. Using meta-analysis, Rotundo and Westgate (2009) analyzed
the environmental effects on component content (mg per seed)
and component concentration (mg g1 ) of soybean seeds. They
concluded that the increase in protein concentration did not neces-
sarily result from a stimulation of protein synthesis, but rather from
a concentration effect due to reduced biomass production under
stress. During water or heat stress, the synthesis of all the major
seed components (protein, oil and residuals) was inhibited, but the
accumulation of individual components was not inhibited to the
same extent (Rotundo and Westgate, 2009). The negative impact
273
was more pronounced on oil and residuals than on protein, which
resulted in a signicant increase in protein concentration. In the
case of water stress, the changes in protein concentration depended
on the timing of stress occurrence and the species. Water stress
imposed throughout the plant life cycle of soybean reduced the
absolute production of all seed components to the same extent. As
a consequence, the nal concentrations of protein in the seed were
not affected signicantly (Rotundo and Westgate, 2009). How-
ever, water stress imposed during late reproductive development
decreased the protein accumulation to a smaller extent than oil
and residuals, leading to a signicant increase in the nal protein
concentration of the stress-exposed soybean seeds. In contrast, an
increase in protein concentration was consistently associated with
a decrease in grain yield of wheat when water decit occurred
throughout the growing season (Ozturk and Aydin, 2004; Flagella
et al., 2010).
Other common responses of plants to environmental stresses
such as drought, heat and elevated ozone include accelerated leaf
senescence and shortened grain development (Black et al., 2000;
Dupont and Altenbach, 2003; Barnabas et al., 2008; Booker et al.,
2009). Accelerated leaf senescence leads to nitrogen remobilization
from vegetative tissue, and amino acids derived from the protein
degradation compensate for the stress-induced decrease in grain
lling time and the nitrogen shortage due to a signicant reduc-
tion of nitrogen uptake from soil by plant roots under stressed
conditions (Triboi and Triboi-Blondel, 2002). As a consequence
of this enhanced amino acid remobilization, protein synthesis is
less affected by stresses than other components (Rotundo and
Westgate, 2009). In contrast, synthesis of carbohydrates in seeds
depends primarily on concurrent carbon xation during seed lling
(Yamagata et al., 1987; Gebbing and Schnyder, 1999). In a number
of studies, grain starch concentration under stress conditions was
reduced due to shortened duration of starch accumulation (Guedira
and Paulsen, 2002; Altenbach et al., 2003; Hurkman et al., 2003),
or due to the inhibition of key enzymes involved in starch syn-
thesis (Hawker and Jenner, 1993; Keeling et al., 1993; Hurkman
et al., 2003). Consequently, adverse growing conditions that pro-
mote leaf senescence during grain lling tend to favor protein
deposition over starch accumulation in the grain, because the pro-
duction and translocation of carbohydrates to the grain is more
sensitive to environmental stresses than protein accumulation
(Pleijel et al., 1999; Fernandez-Figares et al., 2000; Triboi and Triboi-
Blondel, 2002; Rharrabti et al., 2003). In other words, in seeds
grown under stress protein tends to be less diluted by carbohydrate
accumulation, resulting in the observed increase of grain protein
concentration.
3. Lipids
Interactions between environmental stresses and the oil con-
centration or quality of crops have mostly been studied in typical
cooking oil crops, or in medicinal plants containing essential oils
as active compounds. The major stress factors investigated were
drought, salinity, and heat (Table 2). The observed effects were
variable and dependent on the stress type, crop species, and exper-
imental conditions, but some general patterns can be established.
Notably, the majority of the studies reported decreases in the lipid
concentration when crops were grown under stress conditions.
This was particularly true for drought stressed crops, in which
almost all studies reported a decrease in the lipid concentration
of the harvested fractions compared to sufciently watered plants
(Table 2). Only a few studies did not detect signicant decreases
in oil concentration due to drought, and only one study reported
an increase in oil concentration in olives (Palese et al., 2010), per-
haps because olive trees were less susceptible to water decit than
274 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286

Table 2 annual crop species. A similar trend towards declining oil con-
Effects of environmental stress factors on the lipid concentration of the harvested
centration was seen under salinity and heat stress, where only a
fractions of crops.
few studies reported increases or no effects on lipid concentrations
Crop species Effect References (Table 2). In contrast, ozone stress seemed to be an exception, as
Drought the available studies reported either no effect, or even an increase
Chamomile, Matricaria Baghalian et al. (2008) in lipid concentration (Table 2).
recutita L. Apart from the effects of environmental stresses on the oil con-
Corn, Zea mays L. Harrigan et al. (2007)
centration of crops, a number of studies reported changes in the
Groundnut, Arachis Dwivedi et al. (1996)
hypogeal L. fatty acid composition of the harvested oil. A general trend was
Lupine, Lupinus ssp. Carvalho et al. (2004) observed towards an increase in the saturation level of the oil frac-
Nigella sativa L./Plantago Bannayan et al. (2008) tion due to environmental stress, i.e. a decrease in the proportion
ovate Forsk.
of (poly)unsaturated fatty acids and an increase in the proportion
Olive, Olea europea L. Palese et al. (2010)
Rapeseed, Brassica napus L. Bouchereau et al. (1996), of saturated fatty acids. Polyunsaturated fatty acids (PUFA), such as
Pritchard et al. (2000), linoleic acid, are of particular importance due to their health bene-
Aslam et al. (2009), and ts in human diets. A signicant drop in the proportion of PUFA was
Tesfamariam et al. (2010) seen under drought stress in the oil fractions of sunower (Flagella
Safower, Carthamus Movahhedy-Dehnavy et al.
et al., 2002), groundnut (Dwivedi et al., 1996) and sage (Bettaieb et
tinctorius L. (2009) and Ashra and
Razmjoo (2010) al., 2009); under salt stress in sunower (Di Caterina et al., 2007),
Sage, Salvia ofcinalis L. Bettaieb et al. (2009) olive (Chartzoulakis, 2005), cotton (Ahmad et al., 2007), sage (Ben
Soybean, Glycine max (L.) Dornbos and Mullen (1992) Taarit et al., 2010), and coriander (Neffati and Marzouk, 2008); and
Merr. and Foroud et al. (1993)
under heat stress in soy (Dornbos and Mullen, 1992). In many cases,
Sunower, Helianus annuus Flagella et al. (2002)
L.
these decreases in PUFA (especially linoleic acid) were consistent
Wheat, Triticum aestivum L. Zhao et al. (2009) with increases in the proportion of oleic acid (Bouchereau et al.,
Heat 1996; Dwivedi et al., 1996; Flagella et al., 2002; Ahmad et al., 2007;
Creeping bentgrass, Larkindale and Huang Di Caterina et al., 2007; Ben Taarit et al., 2010).
Agrostis stolonifera L. (2004)
Many authors attributed such changes in fatty acid composition
Jojoba, Simmondsia Guillotsalomon et al.
chinensis (Link) (1991) to the activity of enzymes involved in lipid synthesis and conver-
C.K.Schneid sion (e.g. Bouchereau et al., 1996; Flagella et al., 2002). The initial
Rapeseed, Brassica napus L. Aksouh et al. (2001) steps of fatty acid synthesis in oil seeds are localized in the plastids.
Soybean, Glycine max (L.) Dornbos and Mullen (1992)
Oleic acid (a mono-unsaturated fatty acid) is the main product of
Merr.
Spinach, Spinacia olerracea Guillotsalomon et al.
plastidal lipid synthesis, and is subsequently exported to the cytol-
L. (1991) sol. Cytololic desaturation of oleic acid to form PUFA (i.e. linoleic
Salinity acid) is mediated by the enzyme oleate desaturase. The activity
Basil, Ocimum basilicum L. Said-Al Ahl et al. (2010) of this enzyme was put forward as an explanation for shifts in
Chamomile, Matricaria Baghalian et al. (2008)
oleic acid/linoleic acid ratio in several crops under various types
recutita L.
Coriander, Coriandrum Neffati and Marzouk (2008) of stress, including salinity, drought, and heat (Bouchereau et al.,
sativum L. 1996; Flagella et al., 2002; Di Caterina et al., 2007; Hernandez
Cotton, Gossypium Ahmad et al. (2007) et al., 2009). The temperature dependence of this enzyme was
hirsutum L.
proven in several experimental studies (Garcia-Diaz et al., 2002;
Jojoba, Simmondsia Chretien et al. (1992)
chinensis (Link)
Esteban et al., 2004; Rolletschek et al., 2007). In sunower, oleate
Schneider desaturase activity peaked at 20 C and declined at higher tem-
Moringa oleifera Lam. Anwar et al. (2006) peratures, while in safower it was somewhat more heat stable
Olive, Olea europea L. Wiesman et al. (2004), and maintained its maximum activity up to 30 C. Two factors can
Chartzoulakis (2005), Ben
explain this temperature-dependent decline in enzyme activity
Ahmed et al. (2009),
Melgar et al. (2009), (Rolletschek et al., 2007): (i) the heat stability of the enzyme, and
Stefanoudaki et al. (2009), (ii) the effects of temperature on the internal oxygen concentra-
and Mousa (2010) tion of seeds, which is a key regulator of oleate desaturase activity.
Peppermint, Tabatabaie and Nazari Apart from enzymatic desaturation of fatty acids, transport of fatty
Mentha piperita L. (2007)
Rapeseed, Brassica napus L. Boem et al. (1994), Boem
acids from the plastids to the cytolol was suggested as a mech-
et al. (1997), Steppuhn anism that might be affected by environmental stresses (Flagella
et al. (2010), Qasim et al. et al., 2002).
(2003), and Ahmadi and
Ardekani (2006)
Safower, Carthamus Bassil and Kaffka (2002)
tinctorius L. 4. Non-structural carbohydrates
Sage, Salvia ofcinalis L. Ben Taarit et al. (2010)
Stock, Matthiola incana (L.) Heuer et al. (2005) With respect to the impact of environmental stresses on
W.T.Aiton non-structural carbohydrates, two components are of interest as
Sunower, Helianus annuus Francois (1996) and Di
L. Caterina et al. (2007)
determinants of crop quality: (i) the starch concentration of starchy
Sweet fennel, Foeniculim Ashraf and Akhtar (2004) staple crops, and (ii) the sugar concentration or composition in
vulgare (L.) Mill. sugar crops, fruits or vegetables. Moreover, carbohydrates are
Tropospheric ozone important because they are substrates in food processing, for exam-
Bean, Phaseolus vulgaris L. Iriti et al. (2009)
ple in the fermentation of wine.
Soy bean, Glycine max (L.) Mulchi et al. (1988) and
Merr. Heagle et al. (1998) In cereal grains, exposure to environmental stresses during the
grain lling stage was, in most cases, associated with a reduction in
() Stress increased lipid concentration; () stress decreased lipid concentration; ()
stress had no clear effect on lipid concentration.
the starch accumulation. For example, this tendency was observed
in corn under heat, drought, and UV stress (Commuri and Jones,
 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286
1999; Gao et al., 2004; Harrigan et al., 2007), in wheat under heat
stress (Labuschagne et al., 2009), in barley under drought and heat
stress (Savin et al., 1996; Wallwork et al., 1998; Savin and Nicolas,
1999), and in rice under salinity stress (Siscarlee et al., 1990). High
ozone stress seemed to be an exception, as it did not affect the
starch concentration in wheat grains in studies by Fuhrer et al.
(1992) and Scotti et al. (1994). The major mechanism contribut-
ing to these reductions in the starch concentration appeared to
be the inhibition of the enzyme starch synthase in the maturing
grain, resulting in impeded conversion of sugars to starch. Nega-
tive effects of environmental stresses on the activity of this enzyme
were reported in several cereal crops, for example, in wheat under
heat stress (Rijven, 1986; Keeling et al., 1993), in maize under heat
stress (Singletary et al., 1993), or in rice under salinity stress (Khan
and Abdullah, 2003). On the transcript level, reduced expression of
starch synthase genes were detected both in the rice and the wheat
grain under heat stress (Szucs et al., 2010; Yamakawa and Hakata,
2010).
Compared to cereal grains, more variable results were seen
in a number of studies on starchy tuber crops. Enhanced starch
concentration was observed in potato tubers under salinity (Silva
et al., 2001), and heat stress (Debon et al., 1998). In contrast
Bethke et al. (2009) explained increases in the sucrose concentra-
tion with reduced starch formation under drought stress, and no
effect was seen in potato plants exposed to high ozone concentra-
tions (Donnelly et al., 2001). In cassava tubers, a decrease in the
starch concentration was noted under drought stress (Santisopasri
et al., 2001). Two different experiments with sweet potato (Ipomea
batatas) exposed to heat stress reported both a decrease in the
starch concentration (Kano and Mano, 2002), and an increase that
was consistent with increasing amylose content (Noda et al., 2001).
These seemingly contradictory results obtained from experiments
with starchy tubers could be explained by four factors: (i) analytical
artifacts could play a role, as some studies (Debon et al., 1998; Silva
et al., 2001) reported data on fresh weight basis, thus neglecting
stress effects on water concentration of tubers. It is thus possi-
ble that the increases in the starch concentration reported in these
studies were due to a concentration effect caused by lower water
content. (ii) Differences in experimental protocols limit the com-
parability of studies. For example, high temperatures were applied
throughout the day in a study by Noda et al. (2001), while they
were applied only at night in the study by Kano and Mano (2002).
(iii) Enzymatic conversion of sugars to starch, and vice versa, in
tuber crops appears to be strongly affected by environmental fac-
tors in an inconsistent manner. For example, Debon et al. (1998)
observed reduced starch synthase activity in potato tubers due to
heat stress. In contrast, the enzyme -amylase, which decomposes
starch to maltose, tends to be highly activated in photosynthetic tis-
sue during heat stress (Kaplan and Guy, 2004; Kaplan et al., 2006),
but was activated by low temperatures in potato tubers (Nielsen
et al., 1997). (iv) Genotypic differences may lead to contrasting
responses to stress. For example, Watkinson et al. (2008) exam-
ined the expression of starch synthesis and decomposing genes in
drought stressed potato tubers and found inconsistent regulation
that varied between genotypes.
Interestingly, environmental factors also affected the amy-
lose/amylopectin ratio of cereal grain and tuber starches. Water
stress during the grain lling stage decreased the amylose compo-
nent of wheat starch (Singh et al., 2008). This was explained with
the inhibition of granule bound starch synthase (GBSS) activity, an
enzyme involved in amylose synthesis. In contrast, high tempera-
tures led to increases in the proportion of amylose in wheat (Tester
et al., 1995), potatoes (Debon et al., 1998), and sweet potatoes (Noda
et al., 2001). These changes in starch composition due to heat might
also be attributed to altered activity of the enzyme GBSS (Noda et al.,
2001).
275
Sugar concentration showed inconsistent responses to environ-
mental stresses. Increases in sugar concentration of sugar beets
(Beta vulgaris) were noted under drought (Kenter and Hoffmann,
2002; Bloch and Hoffmann, 2004) and salinity stress (Hajiboland
et al., 2009), while another study reported a decrease under drought
stress (Tsialtas et al., 2009). Drought and salinity stress tended
to increase the sugar concentration of fruits and vegetables, as
shown in studies on tomatoes (Petersen et al., 1998; Zushi and
Matsuzoe, 1998; Veit-Kohler et al., 1999; Fernandez-Garcia et al.,
2004; Sgherri et al., 2008; Wu and Kubota, 2008), cucumbers
(Huang et al., 2009) or grapes (Deluc et al., 2009). However, these
increases in sugars concentrations could be due to concentration
effects, as data were reported on a fresh weight basis. In contrast,
experiments on heat stress detected decreases in the sugar concen-
tration of tomatoes (Rosales et al., 2007) or kiwi fruits (Richardson
et al., 2004). The latter study concluded that carbohydrates in heat
stressed kiwi plants were reduced in the fruits and storage tissue
in favor of vegetative tissues to support current growth.
5. Minerals
A large number of studies have investigated interactions
between soil mineral deciencies or toxicities and the mineral
concentration of the edible parts of crops. These interrelations
are well documented and have been extensively discussed and
reviewed elsewhere (Grattan and Grieve, 1999; Wissuwa et al.,
2008; Cakmak, 2008). However, few studies have dealt with the
effects of non-mineral stresses on the mineral concentration of
crops, while those conducted reported no clear trend. In wheat
grown under elevated ozone (Fuhrer et al., 1990; Pleijel et al.,
2006) higher concentrations of several minerals in grains under ele-
vated ozone were possibly attributed to concentration effects (i.e.
reduced grain mass caused by ozone led to the observed increase in
mineral concentrations). Pleijel and Danielsson (2009), found that
zinc in wheat grains showed high correlation with protein concen-
tration when plants were exposed to ozone alone or in combination
with elevated carbon dioxide. Unlike ozone, carbon dioxide tended
to stimulate yields and lead to a dilution of protein and zinc con-
centrations. In the case of corn, ozone stress had minor effects on
macronutrient concentration of K, P and Mg, but the levels of the
micronutrients Zn, Fe and Cu in corn grains increased with increas-
ing ozone concentration (Garcia et al., 1983). Piikki et al. (2007)
found that ozone exposure of potato plants was associated with
increased concentrations of K and Mg in the tubers, while the con-
centration of Ca was unaffected. Two possible mechanisms were
suggested to explain this nding: (i) ozone impeded the carbon
assimilation and thus the total biomass accumulation was reduced
to a larger extent than mineral uptake by the plants; and (ii) plants
exposed to ozone had more wilted haulm at harvest and mobile
mineral elements had been reallocated to the tubers to a larger
extent. Increased carbon dioxide concentrations that were also
investigated in this study did not signicantly affect the concentra-
tions of the investigated macronutrients per se, but did generally
prevent the ozone induced increases in nutrient concentrations.
Decreased concentrations of K, Ca, Mg, Zn in mustard seeds exposed
to ozone stress were observed by Singh et al. (2009), but the authors
did not give an explanation for this phenomenon. However, they
found that the application of 1.5 times the recommended NPK
fertilizer provided protection against yield loss and seed mineral
concentration decrease due to ambient O3 .
Since water plays a signicant role in mineral mobilization,
water deciency may reduce the uptake of Fe, Zn and Cu from
the soil, resulting in decreased concentration of these minerals in
corn grains (Oktem, 2008). However, Ti et al. (2010) found that
severe water decit treatment signicantly increased Ca (28%),
276 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286

Mg (11%), Cu (18%), and Zn (33%) concentrations in maize grains Table 3

Effect of environmental stress factors on phenolics concentration of the harvested
when compared to a fully watered treatment. In contrast, signif-
fractions of crops.
icant decreases in P (17%) and K (17%) concentration in the grain
were seen in the same treatment, which the authors found to be Crop species Effect References
caused by lower soil moisture leading to decreased availability of Drought
these minerals. Conversely, they assumed that soil drought stress Broccoli, Brassica oleracea L. Robbins et al. (2005)
improved routes and/or transport mechanisms for Ca, Mg, Cu, and Grapes, Vitis vinifera L. Esteban et al. (2001) and
Deluc et al. (2009)
Zn, leading to increased grain concentrations in these elements.
Hypericum brasiliense Choisy De Abreu and Mazzafera
Salt stress caused a signicant increase of Na and Cl ions in fruits (2005)
and vegetables (Grattan and Grieve, 1999; De Pascale and Barbieri, Olive, Olea europea L. Palese et al. (2010)
2000; De Pascale et al., 2005; Keutgen and Pawelzik, 2008, 2009), Potato, Solanum tuberosum L. Andre et al. (2009)
Rapeseed, Brassica napus L. Bouchereau et al. (1996)
which is to be expected as these are salt components. As a conse-
Tea, Camellia sinensis (L.) Kuntze Cheruiyot et al. (2007)
quence of ion imbalances caused by Na and Cl accumulation, both Heat
decreases (Gomez et al., 1996; Hasegawa et al., 2000) and increases Hypericum brasiliense De Abreu and Mazzafera
(De Pascale et al., 2005; Keutgen and Pawelzik, 2009) in K con- (2005)
centration were seen in salt stressed plant organs. The decrease in Lettuce, Lactuca sativa L. Oh et al. (2009a)
Potatoe, Solanum tuberosum L. Reyes and
K concentration of plants exposed to NaCl might be related with
Cisneros-Zevallos (2003)
ion competition, implying that excessive Na absorption resulted in Spearmint, Mentha spicata L. Fletcher et al. (2005)
reduced K uptake. In contrast, the accumulation of enhanced levels Spinach, Spinacia oleracea L. Howard et al. (2002)
of K under elevated NaCl might represent a tolerance mechanism, Salinity
Broccoli, Brassica oleracea L. Lopez-Berenguer et al.
as tolerant crop genotypes often have the ability to maintain high
(2009)
K:Na ratios under salt stress (Munns et al., 2006), and the osmotic Lettuce, Lactuca sativa L. Kim et al. (2008a)
function of K might also prevent Na inux into roots and shoots Olive, Olea europea L. Wiesman et al. (2004),
(Jacoby, 1999). Chartzoulakis (2005), Ben
Ahmed et al. (2009), and
Stefanoudaki et al. (2009)
Pepper, Capsicum annuum L. Navarro et al. (2006)
6. Antioxidants Raspberry, Rubus idaeus L. Neocleous and Vasilakakis
(2008)
Due to the outstanding importance of oxidative stress toler- Strawberry, Fragaria ananassa Keutgen and Pawelzik
(2007)
ance in stress physiology (Apel and Hirt, 2004), an overwhelming
Tropospheric ozone
amount of studies have evaluated the response of crops antioxi- Bahiagrass, Paspalum notatum Flugge Muntifering et al. (2000)
dant systems to environmental stresses. However, most of these Bean, Phaseolus vulgaris L. Iriti et al. (2009)
studies dealt with the photosynthetic tissue during the seedling or Clover, Trifolium pratense L. Muntifering et al. (2006a)
early vegetative growth stages of crops, because these are critical and Saviranta et al. (2010)
Lespedeza cuneata Powell et al. (2003)
phases in plant survival and yield formation. Such reports are not
Rice straw, Oryza sativa L. Frei et al. (2010a, 2011)
considered in this review. UV/Light
In contrast, only a limited number of studies reported data on Apple, Malus domestica Borkh. Lancaster et al., 2000;
the antioxidant concentrations in the harvested fractions of crops Merzlyak et al., 2002
Grape, Vitis vinifera L. Douillet-Breuil et al. (1999)
as inuenced by environmental stresses. Such data were reported
and Adrian et al. (2000)
for fruits and vegetables, in which the antioxidant capacity is a Lettuce, Lactuca sativa L. Li and Kubota (2009) and
major factor determining the quality and health effects in human Oh et al. (2009a)
diets. A range of antioxidant components was investigated, includ- Potato, Solanum tuberosum L. Reyes and
ing the hydrophilic classes of phenolics and ascorbate, as well as Cisneros-Zevallos (2003)
Spinach, Spinacia oleracea L. Howard et al. (2002)
the lipophilic classes of carotenoids and tocopherols (Tables 36). A
Tomato, Solanum lycopersicum L. Luthria et al. (2006)
general trend was observed towards an increase in the antioxidant
() Stress increased phenolics concentration; () stress decreased phenolics con-
concentration in crops exposed to stress, but there were exceptions.
centration; () stress had no clear effect on phenolics concentration.
Notably, all studies dealing with high-UV stress found an increase
in antioxidants, and especially phenolics concentration in various
crops (Tables 36). phenolics) may occur in species containing a large fraction of cer-
Phenolics were the most extensively studied antioxidant com- tain phenolic compounds that are degraded under environmental
pounds: a majority of twenty-ve studies reported increases in stress. For example, rosmarinic acid contained in spearmint was
phenolic concentration in the harvested fractions of a variety of shown to be temperature sensitive and strongly degraded under
crops, while only four studies found decreases, and no clear effects heat stress (Fletcher et al., 2005). A further mechanism that could
were noted in an additional ten studies (Table 3). Plant phenolics lead to the reduction in phenolics concentration in older plant
constitute an extremely diverse class of compounds that include tissue (such as forage) is their conversion to more complex phenyl-
phenolic acids, avonoids, anthocyanins, tannins, stilbenes and lig- propanoids, such as lignin and other structural polyphenolics, that
nans (Dai and Mumper, 2010). They all originate from the central are not captured in simple phenolics assays (Muntifering et al.,
phenylpropanoid biosynthetic pathway (Vogt, 2010), and many of 2000; Powell et al., 2003).
them are essential in plants responses to environmental stresses Compared to phenolics, fewer studies have investigated the
due to their antioxidant properties. Therefore, key enzymes of the effects of stress on ascorbate (vitamin C) concentration in harvested
phenylpropanoid pathway, such as phenylalanine ammonia lyase, products, and the results were more controversial (Table 4). Most
are strongly stimulated by diverse environmental stresses as shown studies dealing with tomatoes reported increases in ascorbate con-
in numerous experiments (Kangasjrvi et al., 1994; Guo et al., 2008; centration under drought (Zushi and Matsuzoe, 1998; Veit-Kohler
Oh et al., 2009b; Frei et al., 2010b), and in most cases lead to the et al., 1999; Favati et al., 2009) and salinity (Petersen et al., 1998;
accumulation of phenolics in agricultural products produced under Serio et al., 2004; Sgherri et al., 2007, 2008; Kim et al., 2008b); but
stress conditions (Table 3). However, exceptions (i.e. a decrease in the opposite was reported by De Pascale et al. (2007) under salinity,
 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286 277

Table 4 Table 6
Effect of environmental stress factors on ascorbate concentration of the harvested Effect of environmental stress factors on carotenoid concentration of the harvested
fractions of crops. fractions of crops.

Crop species Effect References Crop species Effect References

Drought Drought
Carrot, Daucus Sorensen et al. (1997) Carrot, Daucus carota L. Sorensen et al. (1997)
carota L. Pepper, Capsicum annuum L. Marin et al. (2009)
Pepper, Capsicum Marin et al. (2009) Potato, Solanum tuberosum L. Andre et al. (2009)
annuum L. Rosemary, Rosmarinus ofcinalis L. Munne-Bosch and Alegre
Potato, Solanum Andre et al. (2009) (2000)
tuberosum L. Sweet potatoe, Ipomoea batatas (L.) Lam. Rautenbach et al. (2010)
Sweet potatoe, Rautenbach et al. (2010) Tomato, Solanum lycopersicum L. Zushi and Matsuzoe (1998)
Ipomoea batatas and Favati et al. (2009)
(L.) Lam. Heat
Tomato, Solanum Zushi and Matsuzoe Lettuce, Lactuca sativa L. Zhou et al. (2009)
lycopersicum L. (1998), Veit-Kohler et al. Tomato, Solanum lycopersicum L. Rosales et al. (2006)
(1999), and Favati et al. Salinity
(2009) Lettuce, Lactuca sativa L. Kim et al. (2008a)
Heat Pepper, Capsicum annuum L. Navarro et al. (2006)
Kiwi, Actinidia Richardson et al. (2004) Tomato, Solanum lycopersicum L. Petersen et al. (1998), De
deliciosa (A. Pascale et al. (2001),
Chev.) C.F. Liang Fernandez-Garcia et al.
& A.R. Ferguson (2004), Serio et al. (2004),
Lettuce, Lactuca Oh et al. (2009a) and Zhou Krauss et al. (2006), De
sativa L. et al. (2009) Pascale et al. (2007),
Tomato, Solanum Rosales et al. (2006) Fanasca et al. (2007), Kim
lycopersicum L. et al. (2008b), and Wu and
Salinity Kubota (2008)
Broccoli, Brassica Lopez-Berenguer et al. UV/light
oleracea L. (2009) Apple, Malus domestica Borkh. Merzlyak et al. (2002)
Cucumber, Cucumis Huang et al. (2009) Lettuce, Lactuca sativa L. Li and Kubota (2009) and
sativus L. Zhou et al. (2009)
Pepper, Capsicum Navarro et al. (2006) Tomato, Solanum lycopersicum L. Rosales et al. (2006) and
annuum L. Becatti et al. (2009)
Raspberry, Rubus Neocleous and Vasilakakis
() Stress increased carotenoid concentration; () stress decreased carotenoid con-
idaeus L. (2008)
centration; () stress had no clear effect on carotenoid concentration.
Strawberry, Fra- Keutgen and Pawelzik
garia ananassa (2007)
Tomato, Solanum Petersen et al. (1998),
lycopersicum L. Fernandez-Garcia et al. and Rosales et al. (2006) under light and heat stress. Furthermore,
(2004), Serio et al. (2004),
increases in ascorbate concentration were reported in lettuce under
Krauss et al. (2006), De
Pascale et al. (2007), various types of stress (Oh et al., 2009a; Zhou et al., 2009), while
Sgherri et al. (2007), Kim drastic decreases were observed under heat stress in the kiwi fruit
et al. (2008b), and Sgherri (Richardson et al., 2004), and under salinity stress in strawberries
et al. (2008)
(Keutgen and Pawelzik, 2007).
UV/light
Apple, Malus Li et al. (2009)
These seemingly contradictory results may be explained by the
domestica Borkh. complexity of the ascorbate metabolism in plants. Generally, the
Tomato, Solanum Rosales et al. (2006) ascorbic acid level of plants is regulated by two processes: (i)
lycopersicum L. de novo synthesis of ascorbate from its precursor glucose-6-P via
() Stress increased ascorbate concentration; () stress decreased ascorbate concen- the intermediates GDP-mannose and l-galactose (Smirnoff et al.,
tration; () stress had no clear effect on ascorbate concentration. 2001); and, (ii) the recycling of ascorbate via the enzymatic network
of the ascorbate-glutathione cycle that restores the reduced form
Table 5 from the oxidized form dehydroascorbate (reviewed by Noctor and
Effect of environmental stress factors on tocopherol concentration of the harvested Foyer, 1998). Both processes have been put forward in explaining
fractions of crops.
shifts in the ascorbate concentration due to stress treatments. For
Crop species Effect References example, Oh et al. (2009a) explained increases in ascorbate concen-
Drought tration in lettuce under relatively mild stress treatments (heat, UV)
Rosemary, Rosmarinus ofcinalis L. Munne-Bosch and Alegre with enhanced expression of an ascorbate biosynthetic gene encod-
(2000) ing l-galactose dehydrogenase. Similarly, high light intensity led to
Soy bean, Glycine max (L.) Merr. Britz and Kremer (2002)
an increased expression of ascorbate biosynthetic genes in apple
Heat
Lettuce, Lactuca sativa L. Oh et al. (2009a) peels, resulting in an enhanced ascorbate concentration (Li et al.,
Soy bean, Glycine max (L.) Merr. Britz and Kremer (2002) 2009). Sgherri et al. (2007, 2008) explained an increase in ascorbate
Salinity concentration in tomatoes under salinity stress with both de novo
Cottonseed, Gossypium hirsutum L. Ahmad et al. (2007) synthesis increasing the total ascorbate pool, and enhanced ascor-
Olive, Olea europea L. Wiesman et al. (2004)
Tomato, Solanum lycopersicum L. Serio et al. (2004), Sgherri
bate recycling via recycling enzymes combined with the reducing
et al. (2007), and Sgherri compound dihydrolipoic acid. In contrast, Richardson et al. (2004)
et al. (2008) concluded that a lack of sugar precursors as substrates for ascorbate
UV/light biosynthesis was the cause of a decrease in ascorbate concentration
Lettuce, Lactuca sativa L. Oh et al. (2009a) and Zhou
in kiwi fruits under heat stress. In a study on strawberries exposed
et al. (2009)
to salinity stress, Keutgen and Pawelzik (2007) hypothesized that
() Stress increased tocopherol concentration; () stress decreased tocopherol con- the ascorbate oxidation rate exceeded the capacity of the recycling
centration; () stress had no clear effect on tocopherol concentration.
systems, which led to a subsequent decline of the ascorbate pool in
278 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286

the fruits. This could be an important mechanism prevailing under Table 7

Effect of environmental stress factors on the digestibility of forage crops and crop
acute exposure to very high stress levels.
residues for ruminant herbivores.
A high level of consensus was seen in the stimulating effects of
various types of environmental stresses on the tocopherol (vita- Crop species Effect References
min E) concentration (Table 5), especially -tocopherol, which has Drought
the highest antioxidant capacity among the vitamin E compounds Alfalfa, Medicago sativa L. Halim et al. (1989, 1990),
(Munne-Bosch and Alegre, 2002). Increases in -tocopherol con- Peterson et al. (1992), Petit
et al. (1992), and Deetz
centration were seen in rosemary and soybeans under drought
et al. (1996)
stress (Munne-Bosch and Alegre, 2000; Britz and Kremer, 2002), Barley straw, Hordeum Leinhos and Bergmann
in olives, tomatoes and cottonseed under salinity stress (Serio vulgare L. (1995)
et al., 2004; Wiesman et al., 2004; Ahmad et al., 2007; Sgherri Brachiaria ssp. Guenni et al. (2002)
Clover, Trifolium ssp. Peterson et al. (1992) and
et al., 2007, 2008), and in soybeans under heat stress (Britz
Lee et al. (2007)
and Kremer, 2002). Similar to ascorbate, shifts in -tocopherol Dactylis glomera L./Lolium Jensen et al. (2003)
concentration due to environmental stress were explained with perenne L.
de novo synthesis (Munne-Bosch and Alegre, 2000) or recycling Mixed forage Jensen et al. (2010)
(Munne-Bosch and Alegre, 2000; Sgherri et al., 2007, 2008). In fact, Maize, Zea mays L. Vincent et al. (2005)
Heat
the recycling of the membrane bound antioxidant -tocopherol
Maize, Zea mays L. Mahmood et al. (2010)
is closely associated with the ascorbate cycle (Smirnoff and Timothy, Phleum pratense L. Bertrand et al. (2008)
Wheeler, 2000; Munne-Bosch and Alegre, 2002): membrane radi- Salinity
cals (-chromanoxyl or -tocpheroxyl), which are formed when Mixed forage Robinson et al. (2004) and
Suyama et al. (2007a,
-tocopherol scavenges lipid radicals, are restored back to -
2007b)
tocopherol by ascorbate, which in turn has to be restored by the Russian thistle, Salsola Fowler et al. (1992)
enzymes of the ascorbateglutathione cycle (Noctor and Foyer, iberica Sennen & Paul
1998). It is probably due to this association between ascorbate Ryegrass, Lolium Ben-Ghedalia et al. (2001)
and -tocopherol that the concentrations in these two compounds multiorum Lam.
Tall wheatgrass, Diaz and Grattan (2009)
showed parallel response to environmental stresses in a number of
Thinopyrum ponticum
studies (Sgherri et al., 2007, 2008). (Podp.) Barkworth &
The major carotenoids investigated were -carotene and D.R.Dewey
lycopene. As with the other antioxidant compounds, a tendency Tropospheric ozone
Alfalfa, Medicago sativa L. Muntifering et al. (2006b)
towards increasing concentration was observed, although with
and Lin et al. (2007)
more frequent exceptions (Table 6). Because lycopene is the major Bahiagrass, Paspalum Muntifering et al. (2000)
carotenoid contained in tomatoes, a number of studies investigated notatum Flugge
the effects of environmental stresses on lycopene concentrations in Clover, Trifolium ssp. Sanz et al. (2005),
tomato fruits. The majority of these studies reported increases in Muntifering et al. (2006a),
and Gonzalez-Fernandez
the lycopene concentration under drought (Zushi and Matsuzoe,
et al. (2008)
1998; Favati et al., 2009), salinity (Petersen et al., 1998; De Pascale Highbush blackberry, Ditchkoff et al. (2009)
et al., 2001; Krauss et al., 2006; Kim et al., 2008b; Wu and Kubota, Rubus argutus Link
2008), and light stress (Becatti et al., 2009); however the opposite Lespedeza Powell et al. (2003)
cuneata/Schizachyrum
(i.e. a decrease in lycopene concentrations in tomato) was reported
scoparium
by De Pascale et al. (2007) under salinity stress, and by Rosales et al. Mixed forage Fuhrer et al. (1994)
(2006) under combined heat and light stress. Consistent increases Poa pratensis L. Bender et al. (2006)
in total carotenoid concentration were observed in lettuce under Rice straw, Ozyza sativa L. Frei et al. (2010a, 2011)
salinity (Kim et al., 2008a), heat (Zhou et al., 2009) and light UV/light
Mixed forage Lindroth et al. (2000) and
stress (Li and Kubota, 2009; Zhou et al., 2009). Such increases in
Thines et al. (2008)
carotenoids could be partly explained with a simple concentration
Digestibility was either determined through in vitro incubation trials in rumen uid
effect because stressed crops had lower water content (Krauss et al.,
obtained from stulated animals, or the effects were estimated based on the chem-
2006; Navarro et al., 2006). However, most authors assumed that ical composition; () stress increased the digestibility; () stress decreased the
biochemical factors were responsible for shifts in carotenoid con- digestibility; () stress had no clear effect on the digestibility.
centration due to environmental stresses. Generally the carotenoid
level in plants is regulated by two biochemical factors (Howitt and
Pogson, 2006; Cazzonelli and Pogson, 2010): (i) biosynthesis via the in carotenoid concentration than milder stresses. For example,
common precursor phytoene; and, (ii) carotenoid storage capac- Munne-Bosch and Alegre (2000) argued that drastic drought stress
ity of the plant tissue, i.e. the formation of sequestering structures led to the destruction of carotenoids in rosemary through photoox-
capable of storing carotenoids in the plastids. A number of studies idation, and Dumas et al. (2003) reported a decrease in lycopene
have attributed increases in the carotenoid concentration of crops concentration in tomatoes exposed to excessive UV radiation asso-
to enhanced biosynthesis (Krauss et al., 2006; Kim et al., 2008a; ciated with visible damage.
Becatti et al., 2009). Indeed, it is well established that phytoene
synthase, the rate-limiting enzyme in the carotenoid biosynthesis, 7. Feed value
is stimulated by environmental factors such as light, heat, drought
and salinity (Cazzonelli and Pogson, 2010). In contrast, reductions A number of studies evaluated the inuence of environmen-
in carotenoid levels due to stress, as seen in a number of studies tal stress factors on the feed quality for ruminant herbivores of
(Table 6), are more difcult to explain. Under heat stress, the tem- forage crops and crop residues (Table 7). The most important qual-
perature dependence of carotenoid biosynthesis may play a role. ity criterion was the digestibility for ruminants, which can either
Dumas et al. (2003) reported that lycopene synthesis was strongly be determined by in vitro incubation experiments using rumen
inhibited at temperatures exceeding 30 C. Moreover, an extremely uid obtained from stulated animals, or it can be estimated based
high level of stress is presumably more likely to induce reductions on the chemical composition. Chemical parameters affecting feed
 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286
digestibility include the structural ber fractions (neutral detergent
ber NDF, acid detergent ber ADF, lignin), the mineral concen-
tration and composition, as well as crude protein and phenolics
concentration. Increases in the ber fractions (especially lignin)
and phenolics are generally thought to decrease the digestibility
of plant materials, while increases in protein concentration are
seen as positive (Traxler et al., 1998; Mould, 2003). Notable dif-
ferences were seen in the effects of different types of stress on
feed value. While all studies dealing with ozone reported a reduced
digestibility of plant materials grown under stress, most stud-
ies dealing with drought reported an enhanced feed value, while
mixed or no effects were seen under salinity, heat and UV stress
(Table 7).
The major chemical factor contributing to the reduced
digestibility of plant materials exposed to ozone was an increase
in the lignin concentration, which was observed in Poa praten-
sis (Bender et al., 2006), rice straw (Frei et al., 2010a), clover
(Sanz et al., 2005; Muntifering et al., 2006a; Gonzalez-Fernandez
et al., 2008), and bahiagrass (Muntifering et al., 2000). Lignin is
an indigestible component for ruminant herbivores and inhibits
the degradation of other cell wall fractions, such as cellulose,
by rumen microbes (Van Soest, 2006). It originates from the
phenylpropanoid biosynthetic pathway that is of central impor-
tance in the synthesis of phenolic compounds (Vogt, 2010). This
pathway tends to be induced by ozone as a response to oxida-
tive stress (Kangasjrvi et al., 1994; Frei et al., 2010b), and
has therefore been associated with increases in non-structural
phenolics and lignin concentrations in ozone stressed plant
materials.
In strong contrast to ozone, drought stress improved the feed
value in most studies (Table 7). Enhanced feed value due to
drought was explained with increases in the protein concentra-
tion (Petit et al., 1992; Jensen et al., 2003), a higher leaf/stem
ratio of drought stressed plants (Peterson et al., 1992), and reduc-
tions in the ber concentration, more specically the lignin fraction
(Peterson et al., 1992; Petit et al., 1992; Deetz et al., 1996). A phys-
iological explanation for the latter effect was given by Vincent
et al. (2005), who demonstrated in a proteomic approach that
the level of lignin biosynthetic enzymes (such as phenylalanine
ammonia lyase and others) was decreased in maize plants exposed
to drought stress. In their study, the level of lignin biosynthetic
enzymes was highly correlated with the lignin concentration of
the plant tissue. However, it must be noted that the opposite
was reported by Lee et al. (2007), who determined the level of
lignin biosynthetic enzymes and lignication in drought stressed
clover and found increases in these parameters. It is not clear at
this point whether these apparent contradictions can be explained
by differences between species, or different experimental
conditions.
Contrasting results were seen in the effects of salinity on
the feed value of diverse forage crops. Several studies reported
improved feed value in salt stressed plants due to increases in pro-
tein concentration and decreases in structural ber, particularly
lignin concentration (Fowler et al., 1992; Ben-Ghedalia et al., 2001;
Robinson et al., 2004). However, other authors pointed to the dan-
ger of accumulating high levels of potentially toxic minerals when
plants were irrigated with saline water. In a study on tall wheat-
grass, Diaz and Grattan (2009) warned that tissue concentrations
in boron, molybdenum and sulfur exceed the maximum tolerable
level for long term intake by ruminants. Similarly, Suyama et al.
(2007a, 2007b) pointed to the problem of excess sulfur and sele-
nium concentrations in a number of forage species irrigated with
saline-sodic drainage water.
Few studies have dealt with the effects of heat and high UV radi-
ation on the feed value of crops (Table 7) and the results were
variable (heat) or only very weak (UV). Further studies might be
279
necessary to obtain a more comprehensive understanding of the
inuence of these factors on forage quality.
8. Physical and sensory traits
Besides nutritional factors, physical (seed mass and shape) and
sensory (seed color, texture, avor) factors are important in deter-
mining the market value of food crops. In this review, we mainly
focus on summarizing the effects of environmental stresses on
physical and sensory factors of two important cereals, rice and
wheat. Other crops, such as maize and soybean, are only briey
described since few studies have been conducted to date. For fruits
and vegetable crops, the readers may refer to recent reviews by
Booker et al. (2009), Moretti et al. (2010) and De Orduna (2010).
Environmental stresses had a negative impact on the appear-
ance of many crops. For example, drought stress negatively
inuenced the appearance of soybean seeds (Bandeh and Jalilian,
1997). Physical properties of cereals grains were affected by envi-
ronmental stresses in many ways: soil salinity reduced the grain
weight of rice (Gay et al., 2010); high temperatures decreased the
kernel weight of corn (Wilhelm et al., 1999), barley (Savin et al.,
1996), rice (Tashiro and Wardlaw, 1991; Morita et al., 2005) and
wheat (Stone et al., 1997; Wardlaw et al., 2002); and drought stress
decreased the grain size of wheat (Zhang et al., 1998). Elevated
ozone concentration decreased the grain weight of wheat in open-
top chambers (Pleijel et al., 1991; Fuhrer et al., 1992; Piikki et al.,
2008), while it had little effect on grain mass of rice grown under
fully open-air eld conditions (Shi et al., 2009). Seed weight of soy-
bean was decreased by ozone in both chamber (Morgan et al., 2003)
and open-air eld conditions (Morgan et al., 2006).
Smaller grain size might be partially responsible for concentra-
tion increases in important components determining the sensory
quality of grains. In rice, 2-acetyl-1-pyrolline (2-AP) is consid-
ered the key aroma compound present in almost all aromatic rice
varieties (Pachauri et al., 2010). A signicant negative correlation
between the 1000-grain weight and 2-AP concentration of three
fragrant rice cultivars was found under soil salinity conditions (Gay
et al., 2010). Yoshihashi et al. (2004) found that drought stress
applied during the milky stage of the ripening phase led to an
increase in 2-AP concentration in the grains at harvest. The positive
effect of osmotic stress, i.e. drought and salinity, on the aromatic
quality of rice may be associated with a fact that osmotic stresses
usually induces the accumulation of proline, which is one of the
precursors of 2-AP (Buttery et al., 1983; Maraval et al., 2008).
Unlike most other cereal grains, visual appearance is an impor-
tant quality criterion in rice, because it is consumed as a whole
grain. Among many factors contributing to visual appearance of
rice grains, chalk (an opaque area in the rice grain) is an impor-
tant quality characteristic in rice and occurs most commonly when
grains are exposed to high temperatures during their development
(Tashiro and Wardlaw, 1991; Lisle et al., 2000; Cooper et al., 2008;
Tsukaguchi and Iida, 2008; Ishimaru et al., 2009; Chun et al., 2009).
Chalky grains have a negative impact on the rice quality, not only
due to their undesirable appearance, but also due to their lower
mill yield since chalky grains tend to be weak and break easily.
Moreover, altered cooking and eating quality of chalky rice was
demonstrated by several studies published internationally (Lisle
et al., 2000; Zhong et al., 2005; Chun et al., 2009) and many reports
in Chinese, but there is no consensus on this aspect (Hu et al.,
2003). High-temperature stress during the grain development facil-
itated the formation of chalky grains through the loose packing
of amyloplasts (Ishimaru et al., 2009). The loosely packed struc-
ture and rapid water absorption of chalky grains were associated
with a more short-branched amylopectin structure and lower hard-
ness, and led to shorter cooking time and lower gelatinization
280 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286
temperatures. The longer chewing time observed in chalky grains
was explained with a lower protein concentration in chalky rice.
Moreover, low amylose concentration of chalky rice caused more
soluble solid and higher iodine index during cooking, indicating a
low eating quality (Chun et al., 2009). By using proteomic analy-
sis, Lin et al. (2005) found that the appearance of chalky kernels
was positively correlated with the expression of small heat shock
proteins, which were induced by high temperature. They hypoth-
esized that small heat shock proteins might affect the aggregation
of macromolecules, such as starch grains or storage proteins, dur-
ing grain lling, and change the physicochemical properties of rice
grains.
In wheat, dough strength is an important quality trait,
which is closely associated with our protein concentration and
composition. Under normal growth conditions, there was a pos-
itive relationship between our protein percentages and dough
strength, whether the latter was determined using maximum
resistance, extensilility, mixing time, loaf volume or SDS (sodium
dodecyl sulfate) sedimentation volume as a quality criterion (Stone
et al., 1997). Gliadins and glutenins are two main classes of gluten
storage proteins responsible for dough strength and baking quality.
Gliadins are prolamins inuencing the extensibility and viscosity
of the dough, while glutenins provide for elasticity of the dough.
Both gliadins and glutenins are essential during the baking pro-
cess, giving bread the ability to rise properly and x shape, and they
are responsible for the rmness of the dough in baked bread. The
ratio of gliadins to glutenins and the proportions of large glutenin
polymers are therefore widely used as indicators of dough strength
(Shewry et al., 1992; Tatham et al., 1985). Heat stress had a negative
impact on the dough strength of wheat our (Wrigley et al., 1994;
Stone et al., 1997; Gibson et al., 1998; Wardlaw et al., 2002). The
heat-related weakening in dough strength was associated with a
higher ratio of gliadin to glutenin because gliadin synthesis contin-
ued at a higher rate than glutenin synthesis during a period of heat
stress (Blumenthal et al., 1993). Wardlaw et al. (2002) also found
that the decrease of dough strength due to heat was related to a
decrease in the proportion of the larger molecular size glutenin.
However, short-term water stress during the grain lling stage
caused an improvement in dough strength, which was observed
consistently with an increase in the amount of glutenin macrop-
olymers (Flagella et al., 2010).
9. Discussion
The previous chapters have shown that all quality traits of crops
can be affected by abiotic environmental stresses. These changes in
quality are caused by a mix of physiological reactions in crop plants
that can be very similar in spite of different types of stress (Fig. 1).
We have classed the physiological responses into six categories
(Fig. 1):
(i) Altered gene expression and enzyme activity are extremely
complex reactions. Transcript proling studies in crop plants
demonstrated that the expression of thousands of genes was
affected by stresses such as heat (Frank et al., 2009), drought
(Degenkolbe et al., 2009), ozone (Frei et al., 2010b), salinity
(Walia et al., 2005), and UV-radiation (Pontin et al., 2010).
Similarly, numerous stress reactions were observed in pro-
teomic studies under abiotic environmental stresses (Ahsan
et al., 2010; Gammulla et al., 2010). The responses of genes
and enzymes involved in the biosynthesis of compounds such
as starch, lipids, and antioxidants induces changes in crop qual-
ity as explained in the respective chapters of this review. The
molecular regulation of these processes and their interactions
with stress is a highly innovative eld of research that may ulti-
mately offer great perspectives for the improvement of crop
quality.
(ii) Oxidative stress caused by the accumulation of reactive oxygen
species (ROS) is a common denominator of most environmen-
tal stresses, including those that form part of this review (Apel
and Hirt, 2004; Munns et al., 2006; Kangasjrvi et al., 2005).
It provokes numerous reactions in plants, including responses
of the antioxidant systems that tend to lead to an increase in
antioxidant concentrations in crops (Tables 36). Antioxidant
enzymes such as peroxidases are also involved in other pro-
cesses such as lignin biosynthesis (Vogt, 2010) that may thus be
stimulated by oxidative stress. In addition, ROS have important
signaling functions in plants, and are involved in the induction
of numerous processes such as programmed cell death and leaf
senescence (Van Breusegem and Dat, 2006).
(iii) Accelerated senescence shortens the maturity period of crops
and affects nutrients translocation processes. This phe-
nomenon is observed in crops under many types of stress
including those that are discussed in this review (Egli, 2004;
Lutts et al., 1996; Kakani et al., 2004). As described in the chap-
ter protein accelerated senescence induces the translocation
of nitrogen to the reproductive plant parts, while shortening
the maturation time, which tends to favor protein over starch
deposition in cereal grains.
(iv) Reduced water content in crops is seen most frequently under
those stresses that have direct effects on the water house-
hold, such as drought, heat, and salinity. As a consequence the
concentration of basically all nutrients are increased, which
is particularly relevant in those crops that are usually con-
sumed at a high water concentration, i.e. fruits and vegetables.
Increased levels of sugars and antioxidants were explained
with reduced water content in a number of studies as outlined
in the respective chapters of this review.
(v) Altered mineral uptake and translocation can be a consequence
of stress induced developmental processes such as the reallo-
cation of minerals during maturation, altered water household,
or ion competition in the case of soil salinity. Examples for
these cases are cited in the chapter on minerals.
(vi) Reduced biomass is the most apparent and widely recognized
effect of abiotic environmental stresses on crops, and leads to
quantitative yield losses that are at the focus of many breeding
and agronomic projects for stress environments. But apart from
affecting yields, reduced biomass also affects quality traits via a
concentration effect. As reduced biomass is often the result of
reduced photosynthetic carbon assimilation, the concentration
of all other elements tends to be increased, leading to higher
crude protein and mineral concentrations, as detailed in the
respective chapters.
While the effects of environmental stresses on crop yields are
almost always negative, we have shown that the picture is much
more complex when it comes to crop quality, where both positive
and negative effects occurred. While starch concentration, lipids
(especially PUFA), the feed value, and physical/sensory traits tend
to be decreased, positive effects dominate in terms of protein and
antioxidant concentration, whereas no clear trend can be detected
in sugar and mineral concentration (Fig. 1). It must be noted, how-
ever, that these are just general trends, and that frequent exceptions
do exist (Tables 17).
Based on these ndings, two strategies can be proposed towards
improving the quality of crops grown in stress environments.
(i) Limiting the negative effects on crop quality by reducing the
induction of stress responses. This could be achieved by improv-
ing resistance traits acting upstream of the physiological response
that leads to the deterioration of crop quality. For example, if
drought causes lower oil quality due to enzymatic changes in an
 Y. Wang, M. Frei / Agriculture, Ecosystems and Environment 141 (2011) 271286
Fig. 1. Conceptual model of physiological responses of crops stimulated by ve types of abiotic environmental stress, and their prevalent effects on quality parameters;
note: prevalent effect means that the majority of studies reported changes in a certain parameter as indicated in the model; exceptions occur as detailed in the text and
Tables 17; P/S traits, physical/sensory traits; PUFA, polyunsaturated fatty acids.
oilseed crop, improving the root system of the plant could mitigate
the effect by avoiding drought stress. Even more importantly, the
occurrence of environmental stresses should be counteracted a pri-
ori, for example by appropriate irrigation, or by reducing pollution.
(ii) Exploiting the positive effects of stresses on crop quality without
compromising yield. For example, if salt stress improves the antiox-
idant level of a vegetable crop, mild salinity could be tolerated to
an extent that does not affect yields.
Changes in crop quality due to environmental stresses have
received less attention than yield losses, perhaps because they are
more difcult to detect. They might occur even at stress levels that
do not affect quantitative yields and/or produce visible damage. For
example, Frei et al. (2011) reported deterioration in the feed value
of rice straw even at ozone concentrations that were too low to
affect quantitative yields. Similarly, tomatoes irrigated with saline
water did not exhibit visible damage, but had enhanced antioxidant
concentrations (Sgherri et al., 2007, 2008). Such improvements
or deterioration of quality parameters constitute hidden stress
effects that should be considered in adaptation strategies to envi-
ronmental stresses.
10. Conclusion
It was shown, that both positive and negative effects of envi-
ronmental stresses on crop quality occurred. As these outcomes
depend on numerous confounding physiological, environmental
and experimental factors, literature reports sometimes appear to
be contradictory, even if the same stress type and species were
investigated. In spite of this complexity, some general trends were
observed, such as a stimulation of protein and antioxidant concen-
tration, or lower starch and lipid concentration, and deterioration
of the feed value and physical/sensory traits. This information can
be used in developing strategies to improve the quality of crops
grown in stress environments by stimulating positive stress effects
or limiting negative effects.
Acknowledgements
We thank Rachel S. Schutte for a thorough review of the
manuscript and for helpful input in designing the conceptual
281
model. Moreover, we wish to apologize to those authors whose
work was overlooked or not considered due to lack of access.
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