ADAPTATION OF STRESS IN PLANTS

(Drougth Stress in Rice (Oriza sativa))

de Asis, Haizel Anne T., Abracia, Mary Grace, Zubiaga, Jahzeel G.,
Hular, Katrina, Artieda, Marcky.,Ogatia, Marilyn.

I. Introduction

Rice (Oryza sativa) is the major food crop in Asia and a major source of

food in almost population in the world, but stress conditions such as drought

often cause severe yield loss.

Rice, Oryza sativa belongs to the family Graminae and subfamily

Oryzoidea, the staple food for one third of the world’s population and occupies

almost one-fifth of the total land area covered under cereals. It is grown under

diverse cultural conditions and over wide geographical range. Most of the

world’s rice is cultivated and consumed in Asia, which constitutes more than half

of the global population. It is also the world’s most important food crop and a

primary source of food for more than half the world’s population. Approximately

11% of the world’s arable land is planted annually to rice, and it ranks next to

wheat. The world’s rice production has doubled during last 25 years, largely due

to the use of improved technology such as high yielding varieties and better crop

management practices (Byerlee, 1996). Large areas of rice are grown under

lowland and upland rainfed conditions. These areas respectively occupy 31%

and 11% of the global rice-growing area (IRRI, 2001).

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 Rice (Oryza sativa L.) is a semiaquatic crop plant that requires high levels

of water in the field for proper growth and development (Biswas and Choudhuri

1994). It is also considered a drought-sensitive crop species; however, within this

species, there are considerable varietal differences in sensitivity to this

environmental stress.

Drought stress

Drought stress is one of the main abiotic stresses to rice, and is often at

least partly responsible for low yields (Zhao et.al., 2007). Drought is also the

most important constraint reducing rice yield in rainfed areas.

Early droughts often result in delayed sowing or transplanting. Yield

reductions from early droughts are minimal and result mainly from a reduction in

tiller numbers. Breeding varieties suited to these conditions is an important

element in reducing risk and increasing productivity in drought-prone

environments, but progress in breeding for drought tolerance in rice has been

slow (Venuprasad et al., 2009).

Exposure of plants to certain environmental stresses can lead to the

generation of reactive oxygen species (ROS), including superoxide anion

radicals (O2), hydroxyl radicals (OH), hydrogen peroxide (H2O2) and singlet

oxygen. Injury caused by ROS, known as oxidative stress, is one of the major

damaging factors in plants exposed to environmental stresses such as drought

(Wang et. al., 2004).

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 Drought is the most important limiting factor for crop production and it is

becoming an increasingly severe and it is becoming an increasingly severe

problem in many regions of the world. Drought is a world-spread problem

seriously influencing grain production and quality. In 2006 and 2008, Tao and

Yang et. al, proved that rice as a paddy field crop is particularly susceptible to

water stress. It is also estimated that 50% of the world rice production is affected

more or less by drought. Drought may delay the phenological development of the

rice plant (Inthapan and Fukai, 1988) and affect physiological processes like

transpiration, photosynthesis, respiration and translocation of assimilates to the

grain (Turner, 1986). Plant processes that depend on cell volume enhancement

are particularly sensitive to water deficit. Leaf expansion and leaf gas exchange

rates are two such sensitive processes. At the plant level, reduced leaf area is

probably the obvious mechanism by which plants and crops restrict their water

loss in response to drought (Sadras and Milory, 1996).

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Composition of Drought Stress

The observation that plants under drought stress generate reactive

oxygen species (Price et al., 1989) and the fact that transgenic alfalfa

expressing MnSOD has reduced injury from water deficit (McKersie et al., 1996)

prompted us to study whether rice expressing foreign SOD under the control of

a stress inducible promoter would improve drought tolerance.

The availability of rice genome sequence will permits now the

identification of the function of each of rice genes through functional genomics,

was published in 2002, serving as a gold standard for all future investigation. To

bring some light on functional gene response to drought stress, the present

study was aiming cloning and sequencing to identify candidate gene response

to drought stress condition and determining their relationship with identified

candidate genes, providing some insights into the molecular basis of gene

tolerant to drought in rice.

Bernier et al confirmed in 2008 that drought as an abiotic stress is the

major constraint to rice production in water-limited environments. Early droughts

often result in delayed sowing or transplanting. Yield reductions from early

droughts are minimal and result mainly from a reduction in tiller numbers.

Breeding varieties suited to these conditions is an important element in reducing

risk and increasing productivity in drought-prone environments, but progress in

breeding for drought tolerance in rice has been slow (Venuprasad et al., 2009).
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Cause of Drought Stress in Crop Yield and Plant water use

One of the causes of the drought stress in rice is the soil dehydration.

Generally, the response of plants to soil water deficits can be described as a

sequence of three successive stages of soil dehydration (Fig. 1).

Stage I occurs at high soil moisture when water is still freely available

from the soil and both stomatal conductance and water vapor loss are not limited

by soil water availability. The transpiration rate during this stage is therefore

determined by environmental conditions around the leaves.

Stage II starts when the rate of water uptake from the soil cannot match

the potential transpiration rate. Stomatal conductance declines, limiting the

transpiration rate to a rate similar to that of uptake of soil water, resulting in the

maintenance of the water balance of the plant.

Finally, stage III begins when the stomata are no longer able to limit the

transpiration to that water available from the soil even through stomatal

conductance is at a minimum. At this time the plant must resort to other

mechanisms of drought adaptation if the plant is to survive.

Virtually all major processes contributing to crop yield including leaf

photosynthetic rate, leaf expansion and growth are inhibited late in stage I or in

stage II of soil drying.

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 The focus of stage III is survival and water conservation mechanisms

which will allow the plant to endure under these severe conditions must be used

if available. Plant survival is a critical trait in natural dry-land ecosystems, but for

most agricultural situations, stage III often, but not always has little relevance to

questions about increasing crop yield (IRRI, 2006).

Consequently, the amount of water available up to the end of stage II for

all practical purposes determines the cumulative growth and yield on a particular

soil. Recovery from stage III can only be of relevance to yield performance if

water is added to the system while there is still sufficient time for growth.

Therefore, options involving mechanisms to enhance crop survival thus do not

usually mean any increase in crop yield under severe drought stress conditions.

Increased crop yields and water use efficiency generally require the optimization

of the physiological processes involved in the critical early stages (mainly stage

II) of plant response to soil dehydration (IRRI,2006).

Figure1. Show the sequence of three successive stages of soil dehydration.

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Genetic Composition of Drought Stress

Genetic differences in drought tolerance offer the unique opportunity to

compare changes in metabolic processes in plants under water stress that might

be involved in drought tolerance.

Matsumoto et.al.2005, and Rensink stated that the genotypic variation in

drought tolerance together with the genetic tools available for rice, such as

marker maps, sequence information, and microarrays and the possibility to test

the agronomic relevance of a scientific discovery, make rice a most interesting

model system for research in drought tolerance of grass crops.

Xiong and Zhu in 2002 and Fujita et.al., 2005 claimed that large numbers

of genes are involved in plant responses in drought. These include genes

involved in signal transduction and transcriptional regulation, biosynthesis of

osmotic and other protectors. In addition a study conducted by Lang, Binh, and

Buu in 2010, aimed in cloning and sequencing genes to identify candidate gene

response to drought stress condition and determining their relationship with

identified candidate genes, providing some insights into the molecular basis of

gene tolerant to drought in rice.

Introducing single genes into a plant of interest could not address the fact

that stress tolerance requires the expression of several genes at one time (Pardo

JM., 2010). A solution to this roadblock in genetic modification is to transform

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plants with stress response-related transcription factors, which would be capable

of regulating several downstream genes expressed to protect against drought

(Pardo JM., 2010). Constitutive promoters used in transgenic plants are effective

in producing high expression levels of the gene of interest. However, gene over-

expression may have detrimental physiological and morphological effects on the

plant during normal growth conditions (Pardo JM., 2010; Bhatnagar-Mathur P et

al, 2008). The use of stress-inducible or organ-specific promoters may provide an

important advantage to engineering effective drought tolerant crops. These types

of promoters allow the plant to develop properly during normal conditions.

Stunted growth, decreased fertilization, and germination are a few adverse

effects caused by accumulation of secondary metabolites or other molecules

such as trehalose or H2O2 in transgenic plants constitutively expressing drought

response-associated transcription factors (Bhatnagar-Mathur P et. al., 2008). The

Rab21 and Wsi18 promoters in Oryza sativa have shown to be induced during

water stress signaling and are successful in eliminating the undesirable

phenotypes resulting from the use of constitutive promoters as well as providing

higher stress tolerance (Bhatnagar-Mathur P. et. al., 2008).

Ecological effects of Drought stress in Plant Body

Stress-inducible promoters, Rab21 and Wsi18 in Oryza sativa showed low

basal levels of expression during normal growth conditions and 36- and 65-fold

increases in leaves after exposure to drought conditions. Expression was 500

and 1400 fold increased in flowers (Yi N, Kim YS, et al., 2010). Both promo-ters
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also showed a dramatic increase in root expression levels following drought

stress and could be used in monocot crops like Zea mays, Hordeum vulgare, or

Triticum aestivum (Yi N, Kim YS, et al., 2010). The reproductive stage is

essential for grain production, so promoters that are effective in relieving drought

stress symptoms during this particular stage of plant development would be most

desirable for increasing yield.

Lastly, a former study done by Pastore et.al. during 2007, stated the

possibility of plant mitochondria involvement in cell adaptation to drought stress.

This statement is supported by a research done by Kromer in 1995, which

proves that mitochondria interact with chloroplasts and peroxisomes in the

photorespiratory cycle; this allows excess reducing equivalents produced during

photosynthesis under conditions of restricted Calvin cycle to be eliminated, thus

preventing an over-reduction of the carriers of photosynthetic electron transport.

Interestingly, plant mitochondria may control reactive oxygen species generation

by means of energy-dissipating systems. Therefore, mitochondria may play a

central role in cell adaptation to abiotic stresses, which are known to induce

oxidative stress at cellular level.

The understanding of molecular basis of genes tolerance to drought stress

will help to breed choice rice cultivars under lower water requirements.

Zayandeh-Rood cultivar in which originated from local cultivars, have

higher ability in solute accumulation such as proline and total carbohydrates than

the other new lines. Due to correlation between drought tolerance of Zayandeh-

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Rood and solute accumulation, it may be suggested that the solute accumulation

is one of the mechanisms for drought tolerance in rice.

On the other hand the role of manganese superoxide dismutase

(MnSOD), an important antioxidant enzyme, may play in the drought tolerance of

rice (Wang et. al., 2004).

Conclusion

Increasing amounts of land dedicated to agriculture are experiencing

longer and more extreme drought conditions. Genetic engineering a drought

tolerance trait in crop plants could sustain crop vitality in stressful environments.

The impacts of drought on plant growth and development limit cereal crop

production worldwide. Rice (Oryza sativa) productivity and production is severely

affected due to recurrent droughts in almost all agro ecological zones. With the

initiation of molecular and genomic technologies, emphasis is now placed on

understanding the mechanisms of genetic control of the drought stress response.

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