Effect of gamma (γ) radiation on morphological ... · REVIEW / SYNTHÈSE Effect of gamma (γ)...

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REVIEW / SYNTHÈSE Effect of gamma (γ) radiation on morphological, biochemical, and physiological aspects of plants and plant products Sumira Jan, Talat Parween, T.O. Siddiqi, and X. Mahmooduzzafar Abstract: Research on the basic interaction of radiation with biological systems has contributed to human society through various applications in medicine, agriculture, pharmaceuticals and in other technological developments. In the agricultural sciences and food technology sectors, recent research has elucidated the new potential application of radiation for microbial decontamination due to the inhibitory effect of radiation on microbial infestation. The last few decades have witnessed a large number of pertinent works regarding the utilization of radiation with special interest in g-rays for evolution of superior varieties of agricultural crops of economic importance. In this review, general information will be presented about radiation, such as plant specificity, dose response, beneficial effects, and lethality. A comparison of different studies has clarified how the effects observed after exposure were deeply influenced by several factors, some related to plant characteristics (e.g., spe- cies, cultivar, stage of development, tissue architecture, and genome organization) and some related to radiation features (e. g., quality, dose, duration of exposure). There are many beneficial uses of radiation that offer few risks when properly em- ployed. In this review, we report the main results from studies on the effect of g-irradiations on plants, focusing on meta- bolic alterations, modifications of growth and development, and changes in biochemical pathways especially physiological behaviour. Key words: growth modulation, metabolic alteration, elicitation. Résumé : French French French French French French French French French French French French French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French French French French French French French French French French French French French French Motsclés : ,,. [Traduit par la Rédaction] 1. Introduction With the discovery of X-rays by W.C. Roentgen in 1896, there was hope for employing this new technology as a tool in plant improvement programs. Hugo de Varies in 1901 and 1903 presented (in two volumes) Die mutations theorie in which an integrated concept for the occurrence of mutation was outlined. The first attempts to stimulate plant growth by exposing seeds or growing plants to low doses of ionizing ra- diation or by the use of radioactive fertilizers, dates back to the 1960s (Sax 1963). Low and high-LET (linear energy transfer) ionizing radiations have been used widely in breed- ing programs because they are expected to cause mutations over a wide spectrum (Yamaguchi et al. 2003; Okamura et al. 2003; Zhou et al. 2006). Examples of favourable traits in- duced after exposures are semi-dwarf growth, earlier matur- ity, higher yields, and resistance to diseases (Mei et al. 1994; Received 5 February . Accepted 12 October 2011. Published at www.nrcresearchpress.com/er on XX January 2012. S. Jan, T.O. Siddiqi, and X. Mahmooduzzafar. Department of Botany, Hamdard University, New Delhi-62, India. T. Parween. Department of Biosciences, Jamia Millia Islamia, New Delhi-25, India. Corresponding author: Mahmooduzzafar (e-mail: sumira. [email protected]). 17 Environ. Rev. 20: 1739 (2012) doi:10.1139/A11-021 Published by NRC Research Press PROOF/ÉPREUVE

Transcript of Effect of gamma (γ) radiation on morphological ... · REVIEW / SYNTHÈSE Effect of gamma (γ)...

Page 1: Effect of gamma (γ) radiation on morphological ... · REVIEW / SYNTHÈSE Effect of gamma (γ) radiation on morphological, biochemical, and physiological aspects of plants and plant

REVIEW / SYNTHÈSE

Effect of gamma (γ) radiation on morphological,biochemical, and physiological aspects of plantsand plant products

Sumira Jan, Talat Parween, T.O. Siddiqi, and X. Mahmooduzzafar

Abstract: Research on the basic interaction of radiation with biological systems has contributed to human society throughvarious applications in medicine, agriculture, pharmaceuticals and in other technological developments. In the agriculturalsciences and food technology sectors, recent research has elucidated the new potential application of radiation for microbialdecontamination due to the inhibitory effect of radiation on microbial infestation. The last few decades have witnessed alarge number of pertinent works regarding the utilization of radiation with special interest in g-rays for evolution of superiorvarieties of agricultural crops of economic importance. In this review, general information will be presented about radiation,such as plant specificity, dose response, beneficial effects, and lethality. A comparison of different studies has clarified howthe effects observed after exposure were deeply influenced by several factors, some related to plant characteristics (e.g., spe-cies, cultivar, stage of development, tissue architecture, and genome organization) and some related to radiation features (e.g., quality, dose, duration of exposure). There are many beneficial uses of radiation that offer few risks when properly em-ployed. In this review, we report the main results from studies on the effect of g-irradiations on plants, focusing on meta-bolic alterations, modifications of growth and development, and changes in biochemical pathways especially physiologicalbehaviour.

Key words: growth modulation, metabolic alteration, elicitation.

Résumé : French French French French French French French French French French French French French French FrenchFrench French French French French French French French French French French FrenchFrench French French FrenchFrench French French French French French French French French French French French French French French FrenchFrench French French French French French FrenchFrench French French French French French French French FrenchFrench French French French French French French French French French French French French French French FrenchFrench FrenchFrench French French French French French French French French French French French French FrenchFrench French French French French French French French French French French French FrenchFrench French FrenchFrench French French French French French French French French French French French French French French FrenchFrench French French French French French French FrenchFrench French French French French French French FrenchFrench French French French French French French French French French French French French French French FrenchFrench French FrenchFrench French French French French French French French French French French French FrenchFrench French French French French French French French French French French French French French

Mots‐clés : , , .

[Traduit par la Rédaction]

1. IntroductionWith the discovery of X-rays by W.C. Roentgen in 1896,

there was hope for employing this new technology as a tool

in plant improvement programs. Hugo de Varies in 1901 and1903 presented (in two volumes) Die mutations theorie inwhich an integrated concept for the occurrence of mutationwas outlined. The first attempts to stimulate plant growth byexposing seeds or growing plants to low doses of ionizing ra-diation or by the use of radioactive fertilizers, dates back tothe 1960s (Sax 1963). Low and high-LET (linear energytransfer) ionizing radiations have been used widely in breed-ing programs because they are expected to cause mutationsover a wide spectrum (Yamaguchi et al. 2003; Okamura etal. 2003; Zhou et al. 2006). Examples of favourable traits in-duced after exposures are semi-dwarf growth, earlier matur-ity, higher yields, and resistance to diseases (Mei et al. 1994;

Received 5 February . Accepted 12 October 2011. Published atwww.nrcresearchpress.com/er on XX January 2012.

S. Jan, T.O. Siddiqi, and X. Mahmooduzzafar. Department ofBotany, Hamdard University, New Delhi-62, India.T. Parween. Department of Biosciences, Jamia Millia Islamia,New Delhi-25, India.

Corresponding author: Mahmooduzzafar (e-mail: [email protected]).

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Environ. Rev. 20: 17–39 (2012) doi:10.1139/A11-021 Published by NRC Research Press

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Li et al. 2007). The different results obtained by variousgroups of plants allowed Muller (1927) to first conclude thatthe rate of appearance of sudden heritable change in plantsand animals can be greatly increased by ionizing radiation.Much evidence of modification and injury to plant parts isfound in the extensive literature on the effects of X-ray treat-ment of plants. Johnson (1939) listed the effects produced on70 species of flowering plants. Seedlings from soaked drywheat (Triticum aestivum L.) and barley seeds (Hordeum vul-gare L.) with 1000 and 5000 R reduced all growth respects,but showed increased tillering. Such exposures mainly stimu-lated seed germination, plantlet growth, flowering, plant size,and yield (reviewed by Breslavets 1946). However, it is diffi-cult to compare the current data on plant responses to g-irra-diation as the models and parameters of past experimentsvaried greatly. Thus, the type of irradiation (e.g., acute orchronic), the dose rate or the dose applied, the physiologicalparameters such as the species/variety/cultivar considered, thedevelopmental stage at the time of irradiation and, finally,inter-individual response variations could all different amongstudies (Gunckel et al. 1953; Gunckel 1957; Boyer et al.2009; Kim et al. 2009) and even among genotypes of thesame species (Kwon and Im 1973). Emerging seedlings arethe stage of development most sensitive to radiation. The sur-vival of grain seedlings was determined by using simulatedfallout g exposure. It is well known that the resistance ofplant seeds is stronger than that of animal cells (Casarett1968; Kumagai et al. 2000; Real et al. 2004). Indeed, theseed is a peculiar stage of a plant’s life cycle characterizedby higher resistance to environmental factors because of itsstructural and metabolic traits, when compared with organsat different life stages. There are also differences in sensitiv-ity to irradiation between dry and fresh seeds because of theirwater content and structures that affect the capacity of ions topenetrate and reach the embryo (Yu 2000; Wu and Yu 2001;Qin et al. 2007). Complex tissue organization seems to bemore resistant to the harmful and mutagenic effects of radia-tion because the multicellular status allows cell and tissue re-pair (Chadwick and Leenhouts 1981; Friedberg 1985; Kranzet al. 1994; Huang et al. 1997; Shikazono et al. 2002; Du-rante and Cucinotta 2008). This lack of uniformity is furthercomplicated by the co-existence of experimental data, applieddata from the food industry, and data emerging from acci-dents (e.g., Chernobyl). Further complexity was created byradiation applied in different dosage measurements. To avoidconfusion we have inculcated historical dosimetery (Table 1).Therefore, in the field of plant radiation, doses vary fromonly a few Grays (Gy) up to several hundred Gy in experi-mental irradiations, but can reach kGy in agribusiness or forvarietal selections (e.g., Bhat et al. 2007; Maity et al. 2009).Furthermore, the dose range response is strongly dependenton the species studied. It is difficult to predict a standard re-sponse to g-irradiation in plants; however, some patterns doemerge.g-rays are the most energetic form of electromagnetic radi-

ation and they possess an energy level from 10 keV (kiloelectron volts) to several hundred keV. they are consideredthe most penetrating radiation source compared with othersources such as alpha and beta rays (Kovács and Keresztes2002). g-rays fall into the category of ionizing radiation andinteract with atoms or molecules to produce free radicals in

cells. These radicals can damage or modify important com-ponents of plant cells and have been reported to affect themorphology, anatomy, biochemistry, and physiology of plantsdifferentially, depending on the irradiation level. These ef-fects include changes in the plant cellular structure and me-tabolism e.g., dilation of thylakoid membranes, alteration inphotosynthesis, modulation of the antioxidative system, andaccumulation of phenolic compounds (Kim et al. 2004; Wiet al. 2005). Several cytogenetic and mutational studies havebeen performed with plant systems exposed to ionizing radia-tion (Table 2). The purposes of these studies was both to tryand determine the harmful effects of radiation and to estab-lish the radiation quality and dose range in which benefits,in terms of more productive or in general more suitable plantsystems, could be obtained.

2. Seed germination

The resumption of active growth in the embryo of a seed,as demonstrated by the protrusion of a radicle (embryonicroot axis) is termed germination. Results regarding the effectof radiation exposure on germination are variable, especiallywhen pre-irradiation treatments are involved. In various ex-periments, parameters such as germination rate and per centare reported to increase, decrease, or remain unchanged afterirradiation. Higher exposures were usually inhibitory (Bora1961; Radhadevi and Nayar 1996; Kumari and Singh 1996),whereas some authors refer to the concept of hormesis, thestimulation of different biological processes (e.g., faster ger-mination, increased growth of roots and leaves), that occurswhen seeds are subjected to pre-irradiation with low dosesof a radiation source (Luckey 1980; Bayonove, et al. 1984;Zimmermann et al. 1996; Sparrow 1966; Thapa 1999). Thestimulatory effects of g-rays on germination may be attrib-uted to the activation of RNA synthesis (Kuzin et al. 1975)on castor bean (Ricinus communis L.) or protein synthesis(Kuzin et al. 1976), which occurred during the early stage ofgermination after seeds irradiated with 4 krad (40 Gy). Thiscould be due to the enhanced rate of respiration or auxin me-tabolism in seedlings. The inability of seeds to germinate athigher doses of g-rays has been attributed to several reasons:(i) numerous histological and cytological changes; (ii) disrup-tion and disorganisation of the tunica or seed layer that is di-rectly proportional to the intensity of exposure to g-rays; (iii)impaired mitosis or virtual elimination of cell division in themeristematic zones during germination (Lokesha et al. 1992).The inhibition of seed germination and seedling growth ex-erted by irradiation has often been ascribed to the formationof free radicals in irradiated seeds (Kumagai et al. 2000; Ko-vács and Keresztes 2002). Many studies have been carriedout evidencing the effects of variable doses of g-irradiationlevels on germination, survival rate and lethality (Table 3).

Table 1. Unit equivalences between systems.

SI units Historical dosimetry1 Gray (Gy) 100 R1 Sievert 100 rem = 100 rad10 mGy 1 Roentgen10 mSv 1 rem = 1 rad

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3. Plant growth and developmentGrowth and development in crop plants do not proceed at

constant or fixed rates through time. Plant development is aterm that includes a broad spectrum of processes by whichplant structures originate and mature as growth occurs. Anychange in growth pattern will ultimately affect maturity andyield. g-irradiation of seed has been found to exert pro-nounced effects on plant growth. The exposure to g-irradia-tions can have stimulatory effects on specific morphologicalparameters and can increase the yield of plants in terms ofgrowth (e.g., taller plants), reproductive success (e.g., formedseeds) and ability to withstand water shortage (Dishlers andRashals 1977; Zaka et al. 2002; Maity et al. 2005; Yu et al.2007; Melki and Dahmani 2009). A more detailed descrip-tion of morphological abnormalities were documented byGunckel (1957) and Sparrow (1966). These results allowedseveral authors (Patskevich 1961; Davies and Mackay 1973;Auni et al. 1978) to conclude that the irradiation of seedsprior to sowing held great promise from a viewpoint of itspractical application in agriculture. It was generally observedthat low doses of g-rays stimulated cell division, growth, anddevelopment in various organisms, including animals andplants. This phenomenon, termed “hormesis”, has been dis-cussed at length for various plant species (Luckey 1980; Sa-gan 1987; Korystov and Narimanov 1997). However, the wayradiation influences plant growth and development is still un-known and the available data remains controversial. Indeed,the magnitude of reported hormetic effects of radiation isusually small; being approximately 10% of control valuesand often not providing critical evidence that crop yield issignificantly increased by irradiating seeds (Miller and Miller1987). The morphological, structural, and functional changesdepend on the strength and duration of the g-irradiation doseapplied. The symptoms frequently observed in plants irradi-ated with a low or high dose were enhancement or inhibitionof germination, seedling growth, and other biological re-sponses (Kim et al. 2000; Wi et al. 2005). Although no con-clusive explanations for the stimulation effects of low dose g-irradiation have been available until now, papers support ahypothesis that the low dose irradiation will induce growthstimulation by changing the hormonal signalling network inplant cells or by increasing the antioxidative capacity of cells

to easily overcome daily stress factors such as fluctuation oflight intensity and temperature in growth conditions (Kim etal. 2004; Wi et al. 2007). In contrast, the growth inhibitioninduced by high-dose irradiation has been attributed to thecell cycle arrest at the G2/M phase during somatic cell divi-sion and (or) varying damage to the entire genome (Preussand Britt 2003). The relationship between growth of irradi-ated plants and the dose of g-irradiation has been manifestedby investigating the morphological changes and seedlinggrowth of irradiated plants. Low doses apparently inhibitauxin synthesis while larger doses can destroy auxin activitydirectly. As with other wound responses, irradiated tissuesoften produce endogenous ethylene (Maxie et al. 1966;Dwelle 1975; Chervin et al. 1992; Liu et al. 2008). Manystudies have been carried out to evidence the effects of varia-ble doses of g-irradiation levels on growth and yield attrib-utes (Table 4).Growth inhibition by g-irradiation may be related to auxin

and DNA biogenesis in a relationship as shown indicated inFig. 1. These relationships postulate exclusive possibilities:(1) that DNA is required for and is previously synthesized se-quentially to auxin formation, the radiation block occurringin the formation of nucleic acid; (2) that the primary radia-tion block is in auxin synthesis, the auxin required for theformation of DNA; and (3) that the effect of radiation is onan undefined entity in reaction previous to and essential forboth DNA and auxin synthesis (Lage and Esquibel 1995;Momiyama et al. 1999).g-irradiation had a stimulatory effect on primary branches

and yield attributes, including number of pods per plant,number of flowers per plant, seed index, etc. following lowdoses of g-rays and inhibition of the same attributes at higherrates (Charumathi et al. 1992; Khan et al. 2000; Jan et al.2010). These groups further investigated the effectiveness ofg-irradiation for induction of early flowering in legumes.Yousaf et al. (1991) and Svetleva and Petkova (1992) re-ported inhibitory effects of g-irradiation on seed maturity,flowering, plant height, and seed yield per plant. g-irradiationprolonged the growth period and retarded plant height infrench beans (Phaseolus vulgaris L.). An increased numberof okra fruit per plant and fruit length as a result of g-irradi-ation was recorded by many authors (Dubey et al. 2007; Mis-

Table 2. Effects of exposure to g radiation on chromosome aberrations of different plant parts.

Source of radiation Species Plant part Parameter studied ReferencesX-rays, 1.7 MeV neutrons H. gracilis (Nutt.) Cell suspensions Cell survival Werry and Stoffelsen (1979)g, 16 MeV neutrons N. plumbaginfolia (L.) Protoplasts Cell survival Magnien et al. (1981)g, 40Ar,56Fe Rice (Oryza sativa L.) Seeds Micronuclei and chromosomal

aberrationsMei et al. (1994)

g, H+, He and N+ P. sativum (L.) Seedlings Micronuclei Vasilenko and Sidorenko (1995)g, H+, He, C5+ N. tabacum L. Seeds Chromosomal aberrations Hase et al. (1999)g A. thaliana (L.) Transgenic seed

and seedlingsHomologus recombination Kovalchuk et al. (2000)

g, C5+ N. tabacum (L.) Seeds Chromosomal aberrations inroots of different length

Hase et al. (2002)

g, C5+ N. tabacum (L.) Protoplasts Cell survival Yokota et al. (2003)g60Co,137Cs N. tabacum (L.) Protoplasts Unrepaired dsb Yokota et al. (2007)g T. durum (L.) Seeds Chromosomal aberrations in

the M1 generationCao et al. (2009)

Ne, Si,40Ar A. cepa (L.) Seedlings Micronuclei Takatsuji et al. (2010)

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Table 3. Effects of exposure to g irradiation on seed germination of different plant.

Dosage applied and durationof irradiations Species

Plant part or age /stage ofplant growth for g-irradiation Parameter studied seed germination References

2–50 krad Fennel (Foeniculum vulgare Mill.) Seeds 2 krad resulted 2% stimulation of germinationand 0.7% with 50 krad

Zeid et al. (2001)

0.1–1 kGy, 1.66 kGy h–1 atthe time of irradiation

Lentil (Lens culinaris Medik) Seed Germination percentage was reduced to 40.87%at 0.2 kGy and no germination at 1.0 kGy dose

Chaudhuri (2002)

150–300 Gy Rice (Oryza sativa L.) Seeds Germination decrease from 100% to 97.2% Cheema and Atta (2003)0.085 and 0.15 kGy Sweet potato (Ipomoea batatas L.) Tuber (8.5 and 15 krad) inhibits sprouting Tabares and Perez (2003)1.0–30.0 kR g-radation atthe rate of 2.8 kR/min.

Pinus kesiya and P. wallichiana Seeds Seeds exposed to 30 kR germinated whereas inP. wallichiana 30 kR was lethal and seedgermination was restricted up to 20 kR only

Thapa (2004)

150, 300, 500, 700, 900,1000 Gy; dose rate10 Gy/28.97 s

Maize (Zea mays, Okra(Abelmoschus esculentus andGroundnut (Arachis hypogeal L.)

Seeds 90% germination was achieved at 240 Gy inmaize, 70% at 170 Gy in Okra and 58% at300 Gy in groundnut.

Mokobia and Anomohanran(2005)

300, 400, 500, 600 and800 Gy

Long bean (Vigna sesquipedalis). Seeds 70.56% increase at 400 Gy. At 800 Gy nogermination.

Kon et al. (2007)

300–800 Gy Tomato (Lycopersicon esculentumcv)

Seeds Germination decrease from 88% to 48.70% with800 Gy

Norfadzrin et al. (2007)

300, 400, 500, 600, and800 Gy

Snap bean (Phaseolus vulgaris) Seeds Germination decrease from 75.56% to 51.11%.At 800 Gy failed to germinate

Ellyfa et al. (2007)

20–110 Gy Atropa belladonna Seeds Germination rate increased from 21.18% to93.33% at 110 Gy

Abdelhady et al. (2008)

300, 400, 500, 600, and800 Gy

Chilli (Capsicum annuum) Seeds Decrease in germination 42.22% to 15.56%, seedsirradiated with 800 Gy failed to germinate.

Omar et al. (2008)

100–1200 Gy, 1.66 kGy h–1at the time of irradiation

Chick pea (Cicer arietinum L) Seed Seed germination, 60%–76% increase with 100–500 Gy, 80%–96% decrease with700–1200 Gy

Shah et al. (2008)

50–300 Gy Pumpkin (Cucurbita pepo L.) Seed, Pollen 75.0% increment in germination (at 50 Gy) to63.0% (at 100 Gy)

Kurtar (2009)

20–80 krad g radation at therate of 2.8 kR/min.

Lepidum sativum L. Seeds 92.50%–97.50% increment in germination with8krad

Majeed et al. (2009)

100–400 Gy; dose rate of0.864 kGy/h

Wheat (Triticum aestivum L.) Seeds, Seedlings Germination percentage decrease from 8.8%(100Gy) to 5.5% (400Gy)

Borzouei et al. (2010)

10, 30, 50 kR Corn (Zea mays L.) Seeds 10 kR had 30% germination and the untreatedhad 50% germination.

Itol et al. (unpublished)

300–500 Gy Okra (Abelmoschus esculentusMoench

Seeds Seed germination 87%–91% increase with500 Gy

Hegazi and Hemeldeldin(2010)

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Table 4. Effects of exposure to g irradiation on morphological and yield attributes of different plant species.

Dosage applied and durationof irradiation Species

Plant part or age /stage ofplant growth for g-irradiation Morphological and yield attributes response References

1–20 Gy Chineses Cabbage (Brassicacampestris L.Cv.)

Seeds 45.67% increment in plant height and 14% increase infresh weight with 20Gy.

Kim et al. (1998)

0.5–60 Gy, dose rate 2.94Gy /min

Pea (Pisum sativum) Seeds 20% Decrease in plant height with 6 Gy. Doses be-tween 10 and 40 Gy were lethal.

Zaka et al. (2002)

150–300 Gy Rice (Oryza sativa L.) Seeds 300 Gy causing 50% seedling height reduction andinduction of >71% sterility with the was same in allthe three rice varieties.

Cheema and Atta (2003)

1, 1.5, 2.5, 5, and 10 krad Corn (Zea mays L.) Seeds Grains exposed to 1.5 and 2.5 krad resulted in 84%and 76% decrease in shoot length.

Al Salhi et al. (2004)

2, 4, 8, and 16 Gy Dose rate150 TBq of capacity

Red pepper (Capsicum annuum) Seeds Except for 16 Gy dose seedlings showed 13% to25%, 5% to 20%, or 26% to 44% increase in stemlength, diameter and leaf area, respectively.

Kim et al. (2004)

1.0–30.0 kR g radation atthe rate of 2.8 kR/min.

Pinus kesiya and P. wallichiana Seeds In Pinus .kesiya, root length showed more than 50%inhibtion by g dose 10 kR. While asin P. walli-chaina 10kR proved lethal

Thapa (2004)

2, 4, 8, and 16 Gy, dose rate150 TBq of capacity

Red pepper (Capsicum annuum) Seeds Except for 16 Gy dose seedlings showed 13% to25%, 5% to 20%, or 26% to 44% increase in stemlength, diameter and leaf area, respectively.

Kim et al. (2004)

50–350 Gy; dose rate 120 G/h

Rice (oryza sativa), Mung(Phaseolus vulgaris)

Seeds 50 Gy resulted in 60% increase in seed yield, plantheight in Oryza sativa L., for Phseolus mungo 200Gy resulted in 56% increase in plant height.

Maity et al. (2005)

150, 250, 350, and 450 Gy Wheat (Triticum aestivum L.) Seeds 88.99% callus induction with 150 Gy, 45.98% and28.65% reduction with 350 and 450 Gy. 15.94% in-crement in grain yield and 37.85% decrease ingrainyield with 450 Gy.

Abdelhady and Ali (2006)

10–100 Kr; dose rate 5000ci Co60

Crotalaria saltiana L. Seeds 64.1% and 51.85% increment in plant height with 30Kr and 50 Kr.

Shah et al. (2008)

20–80 krad g radation at therate of 2.8 kR/min.

Lepidum sativum L Seeds Shoot length decreased 76.42% to 28.20%, Majeed et al. (2009)

10–100 Gy, dose rate 1.858Gy/s

Thai Tulip (Curcumaalismatifolia Gagnep)

Seeds Mean survival rate 50% with 20 Gy and 2% survivalat 40 Gy.

Abdullah et al. (2009)

100–400 Gy; dose rate of0.864 kGy/h

Wheat (Triticum aestivum L.). Seeds 80.61%, 62.5%, 31.42% decrease in root length, rootfresh weight and shoot length, respectively.

Borzouei et al. (2010)

200, 700, 1200 Gy Barley (Hordeum vulgare spp.) Seeds 50% reduction in shoot length with 1200 Gy, 29.67%decrease in root number with 1200 Gy.

Nasab et al. (2010)

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hra et al. 2007; Sharma and Mishra 2007). Soehendi et al.(2007) stated that modification obtained by g-irradiation ofmung bean (Vigna radiata L.) leaflet type could affect leafcanopy and alter seed yield. A higher dose of g-irradiation(500 Gy) in this study was less effective than the two otherlower doses. This observation was also stated by (Artık andPekşen 2006), who found a reduction in faba bean (Viciafaba L.) seed yield and harvest index in some varieties whenseeds were treated with relatively low doses (25 and 50 Gy)of g-irradiation. Plant biomass in plants was significantly af-fected by g-irradiation. The biomass was reduced by approx-imately 50% at a dose of 20–60 Gy. This is consistent withthe radiation responses of cereals, in which doses of 20–60Gy reduced yield by 50% depending on the stage of develop-ment at exposure (Fillipas et al. 1992). Carbon partitioning isaltered by increasing the radiation dose because of damage toradiosensitive cells responsible for the transport of carbohy-drates in the phloem (Thiede et al. 1995).

4. Biochemical and physiological effectsg-irradiation can be useful for the alteration of one or a

few physiological characters (Lapins 1983). Photons of g-ir-radiation are powerful enough to be completely democraticwith regard to the molecular species with which they interact(Fig. 1). Based on previous research reports, the total proteinand carbohydrate contents decreased with increasingly higherdosage of g-irradiation caused by higher metabolic activitiesand hydrolyzing enzyme activity in germinating seed (Barroset al. 2002; Maity et al. 2004). Radiation resulted in the in-creased absorption of glucose, pyruvate, and the decreased

absorption of acetate and succinate in carrot (Dacus carrotaL.). Radiation reduced the anabolic utilization of all sub-strates (Bourke et al. 1967). Bourke et al. reported a decreasein all amino acids upon irradiation except for serine and va-line, which were increased at 100 krad (1 Gy). Irradiationwith doses of 15–30 kGy could reduce viscosity of pectinand alginate (Irawati and Pilnik 2001). Subtle differences inthe degradation of oligosaccharides in processed legumeswere reported by (Machaiah and Pednekar 2002) betweencontrol and irradiated dry seeds of Bengal gram (Cicer arie-tinum L.), Horse gram (Macrotyloma uniflorum (Lam.)Verdc.) and cow pea (Vigna unguiculata (L.) Walp.). g-irra-diation breaks the seed protein and produces more aminoacids (Barros et al. 2002; Maity et al. 2004; Kiong et al.2008). This process may also inhibit protein synthesis. Totalproteins and carbohydrates decreased with increasing high g-ray dosage in wheat and rice plants (Hagberg and Persson1968; Inoue et al. 1975). Lester and Whitaker (1996) ob-served the retention of proteins in the plasma membrane ofmuskmelon (Cucumis melo L.) fruit 10 days after treatmentwith 1 kGy irradiation. The effect of g-irradiation on the con-centration of total free amino acid nitrogen of five varietiesof Iraqi dates (Phoenix dactylifera L.) was studied. The mostsensitive amino acids appear to be proline, glutamic acid, as-partic acid, serine, histidine, lysine, and tyrosine, while me-thionine, isoleucine and leucine showed a slight increase intwo varieties (Auda and Al-Wandawi 1980). Extensive re-search has shown that proteins, essential amino acids, miner-als, trace elements and most vitamins do not representsignificant losses during irradiation, even at doses over 10kGy. Pradeep et al. (1993) reported that amino acids like ly-

Enzyme effects

(in vivo)

Enzyme effects

(in vitro)

Biochemical Reactions

Genetic code

Transcription &

translation

Growth, yield,

quality reduction

In vitro changes

In cellular organelles

Metabolic Pathways

CHRONIC EXPOSURE

BIOCHEMICAL LEVEL

Break down of metabolic

Pathways

Changes in cellular

Constituents

Disruption

Cellular organelles

CELLULAR LEVEL

Membrane

Impairment

Cell Changes

Imbalances

Cell Death

Chlorosis

Senescence

Growth, yield

Quality reduction

Reduce vigor, more-

susceptibility

Necrosis

Plant Death

Survival of

Resultant

Individuals

WHOLE PLANT LEVEL

γ-

R

A

Y

S

Fig. 1. Effect of g radiation plant growth, development, and metabolism.

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sine and histidine were resistant to g-irradiation (5 kGy) inGinseng panax leaf tea (herb). Al-Jassir (1992) found thatthe contents of arginine, methionine, lysine, phenylalanine,and leucine of garlic bulbs (Allium sativum L.) increasedslightly. However, reduction in other amino acids in irradi-ated samples also occurred, especially at higher doses.Many studies by (Marchenko et al. 1996; Ussuf et al.

1996) have also shown that irradiated plant cells have a self-protection mechanism. This mechanism works by enhancingthe synthesis of substance groups or enzymes containing sul-fur such as amino acids (cysteine, cystine), glycation, andsuperoxide dismutase, which can protect, remove free radi-cals, and participate in the simultaneous release of protectivesubstances in an organism (Qui et al. 2000). The maximumincrease in sucrose content in both potato tubers and sweetpotato roots was achieved by an irradiation dose of 3 to 4kGy for potatoes (Solanum tuberosum L.) and 0.8 to 2 kGyfor sweet potatoes (Ipomoea batatas L.) Hayashi and Kawa-shima (1982). Low doses of g-rays enhance chlorophyll syn-thesis by increasing the activating enzyme system. Theseresults were almost in agreement with those of other groups(Zeerak et al. 1994; Al-Kobaissi et al. 1997; Gautam et al.1998; Rascio et al. 2001; Osama 2002), who reported thatthe improvement of yield components and chlorophyll pa-rameters in various plants such as tomato (Lycopersicon es-culentum L.), maize (Zea mays L.), rice (Oryza sativa L.),and wheat (Triticum aestivum L.) was induced after variabledoses of g-rays. Soehendi et al. (2007) stated that the modifi-cation obtained by g-irradiation of mung bean leaflet typecould affect leaf canopy and alter seed yield. Thus, modifiedmung bean lines that exhibited greater leaf area per plantwould have enhanced photosynthetic rate and hence gavegreater yield. g-rays were also found to cause modulation inprotein patterns by inducing appearance and (or) disappear-ance of some protein bands (Rashed et al. 1994). Yoko et al.(1996) studied the effect of g-irradiation on the genomicDNA of corn, soybean, and wheat. They concluded that largeDNA strands were broken into small strands at low irradia-tion dose but small and large DNA strands were broken athigher irradiation doses. This observation was also stated byArtık and Pekşen (2006) who found a reduction in faba beanseed yield and harvest index in some varieties when seeds aretreated with relatively low doses 25 and 50 Gy of g-irradia-tion.Higher g-irradiation inhibits chlorophyll synthesis in wheat

(Kovács and Keresztes 2002) and changes in pigmentation(reddening) were observed in the older leaves of Holcus la-natus L. treated with 40, 80, and 160 Gy (Jones et al. 2004).This was demonstrated in etiolated barley and wheat leaves,in potato tubers. In fruits with chloroplast in hypodermis atthe time of harvest (Hardenpont pear) it was found that g-ir-radiation (1 kGy) altered chloroplast structure characteristi-cally (Kovács and Keresztes 1991). Byun et al. (2002)utilized irradiation technology to reduce or eliminate the re-sidual chlorophyll in oil processing without developing lipidperoxidation during irradiation.g-irradiation can be useful for the alteration of physiologi-

cal characteristics (Kiong et al. 2008). The biological effectof g-rays is based on the interaction with atoms or moleculesin the cell, particularly water, to produce free radicals (Ko-vács and Keresztes 2002). These radicals can damage or

modify important components of plant cells and have beenreported to affect differentially the morphology, anatomy, bi-ochemistry, and physiology of plants depending on the radia-tion dose (Ashraf et al. 2003). These effects include changesin the plant cellular structure and metabolism e.g., dilation ofthylakoid membranes, alteration in photosynthesis, modula-tion of the anti-oxidative system, and accumulation of phe-nolic compounds (Kovács and Keresztes 2002; Kim et al.2004; Wi et al. 2007; Ashraf 2009). From the ultra-structuralobservations of the irradiated plant cells, the prominent struc-tural changes of chloroplasts after radiation with 50 Gy re-vealed that chloroplasts were more sensitive to a high doseof g-rays than the other cell organelles. Plastids were affectedby irradiation in two ways: (i) inhibition of senescence and(ii) dedifferentiation into agranal stage (Kim et al. 2004) (Ta-ble 5). The developmental regression of chloroplasts can beassumed primarily from destruction of grana (Kovács andKeresztes 1989). Similar results have been reported to be in-duced by other environmental stress factors such as UV,heavy metals, acidic rain, and high light (Molas 2002; Ga-bara et al. 2003; Quaggiotti et al. 2004). However, the low-dose irradiation did not cause these changes in the ultra-structure of chloroplasts. The irradiation of seeds with highdoses of g-rays disturbs the synthesis of protein, hormonebalance, leaf gas exchange, water exchange, and enzyme ac-tivity (Hameed et al. 2008). The chlorophyll content of g-ir-radiated wheat displayed a gradual decrease at 200 Gy dose(Borzouei et al. 2010). Kiong et al. (2008) reported that thereduction in chlorophyll b is due to a more selective destruc-tion of chlorophyll b biosynthesis or degradation of chloro-phyll b precursors. Furthermore, Kim et al. (2004) haveevaluated the chlorophyll content on irradiated red pepperplants; their results showed that plants exposed at 16 Gymay have ~23% increase in their chlorophyll content that canbe correlated with stimulated growth. Modulation in photo-synthesis in irradiated plants might partly contribute to in-creased growth (Kim et al. 2004; Wi et al. 2007).The effect of presowing g-irradiation treatments on the

growth and respiration of germinating corn was investigated.Exposures of 40 and 80 krads caused marked inhibition inseedling growth and inhibited the rate of respiration, meas-ured as oxygen uptake per seedling (Woodstock and Combs1965). In contrast to radiation chemistry of food proteinsand carbohydrates, where indirect radiation effects mediatedthrough water play a major role, reactions of lipids with reac-tive species of water radiolysis play only minor role in mostsituations, quantitatively at least. The primary effect of anelectron upon fats leads to cation radicals and excised mole-cules. Regardless of localization, the main reactions of thecation radical are deprotonation, followed by dimerization ordisproportionation. Another primary effect is electron attach-ment, which may be followed by dissociation and decarbon-ylation or dimerization (Diehl 1990). Additional reactionscan be initiated from the excited triglyceride molecules. Thisleads to formation of particular free radicals as principal in-termediates, and ultimately to a particular end product. Thelow dose used could have produced its long term effects inpart by means of stimulation of lipid degradation, possiblymediated through the action of free radicals that are knownto be generated after irradiation (Katsaras et al. 1986; Voisineet al. 1991). Increases in irradiation doses to 10 and 15 kGy

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Table 5. Effects of the exposure to g radiation on eco-physiology and biochemistry in different vegetal systems.

Dosage applied and durationof irradiation Species

Plant part or age /stage ofplant growth for g-irradiation Phyiological and biochemical response References

2,4 and 8 kGy; dose rate38.1 kGy/h

Kidney beans (Phaseolusvulgaris L.)

Dry kidney beans 8 kGy increased deamidation by 65.89%, sulfahydryl contentwas reduced to 23.1%

Dogbevi et al. (1999)

3.35, 6.70, 10.05, and 13.40Gy; dose rate of 0.8 mGy/s

Pumpkin (Cucurbitapepo)

Seedlings Relative chlorophyll concentration increased from 6.3% at3.35 Gy to 16.8% at 13.40%

Jovanić and Dramicanin(2003)

Dose range 5–160 Gy; doserate 7.0 Gy/min

Yorkshire fog grass(Holcus lanatus)

Five-leaf growth stage Shoot nitrogen concentration increased to 3.7% at 20 Gy Jones et al. (2004)

0.15 and 0.30 kGy; exposuretime 30 and 55 min, re-spectively

Onion (Allium cepa L.) Bulbs Glucose, fructose, sucrose were decreased to 21.34, 45.65%and 34.8%, respectively

Benkeblia et al. (2004)

2, 4, 6, 8, and 10 k rad; doserate 10 min/k-dose rate

Chamomile (Chamomillarecutita L.)

Seeds 47.2% increment in essential oil, 33.69% increment in carbo-hydrate content with 10k rad and 21.17% increase in sugarcontent

Nassar et al. (2004)

1, 3, 5, 8, 10, 12, and 15 Kr. Sunatoarea (Hypericumperforatum L.)

Leaves Mono saccharide content reduced to12.59% at 5 Kr. At 3 Krpolysaccharide content increased to 23%

Ichim et al. (2005)

50–300 Gy; dose rate0.54 Gy/min

Rocket (Eruca vesicariassp. .sativa)

Seeds At 20 Gy, total sugar increased by 70.3%, total free aminoa-cid by 92.6%, total soluble phenol by 109%. At 200 Gy to-tal sugar decreased by 70.7%, total free aminoacid by 69.5%and soluble phenol by 67.5%

Moussa (2006)

1, 2, 4, 6, 8, and 10 kGy;dose rate 228Gy/min

Soybean seeds (Glycinemax (L.) Merr.)

Seeds 10% increase in total phenols, 21.6% for tannins at 1kGy. At10 kGy increase in phenol 7.6% and tannins 11%

Štajner et al. (2007)

5, 10, 25, and 50 Gy; doserate /min 10 Gy

Paulowinia tomentosa Node explants Chlorophyll a, chlorophyll b and total chlorophyll was re-duced to 34.69%, 33.33% and 35.21% at 25 Gy, respec-tively

Alikamanoglu et al.(2007)

2.5, 5, 7.5, 10, 15, and 30kGy. Dose rate 6.5 kGy/h

Velvet beans (Macunapruriens)

Seeds Protein content increased to 31.06% at 30 kGy, carbohydratecontent increased 2.2% at 15kGy, linoleic acid increased at30 kGy

Bhat et al. (2007)

0, 10, 20, 30, 40, 50, 60,and 70 Gy; dose rate 4.64kGy/h

Orthosiphon stamineus Shoot tips Plantlets irradiated at 10 and 20 Gy exhibited total solubleprotein content of 39.61 and 34.00 mg/g, respectively

Kiong et al. (2008)

2, 4, 8, 10, 12, and 16 kGy;done rate . 0.96 kGy/h

Kalungi (Nigella sativa) Seeds Extraction yields increases were 3.7%, 4.2%, 5.6%, and 9.0%for hexane, acetone, water and methanol extracts. Phenolcontent increase from 3.7 for control to 3.8 mg/g for 16kGy

Khattak et al. (2008)

0, 5, 10, 15, and 20 kGy;dose rate of 2.38 kGy/h

Tea (Camellia sinensis L.) Tea leaves 37.86% of the compounds were stable at all radiation dosesand 47.53% of new compounds were identified after irra-diation

Fanaroi et al. (2009)

300, 400, and 500 Gy Okra (Abelmoschus escu-lentus L.)

Seeds Chlorophyll a, chlorophyll b and total chlorophyll was in-creased to 17.19%, 18.29%, and 14.44% at 500 Gy, respec-tively.

Hegazi and Hamideldin(2010)

100, 200, 300, 400, and 500Gy; dose 10 Gy/min

Soybean (Glycine max L.Merrill)

Seeds Total chlorophyll decrease was 81.36% at 400 Gy and 80.91%at 500 Gy. Increase varied between 4.75% and 87.39% forFe, 24.08% and 163.09% for Cu, and 9.86% and 45.14% forZn with respect to the control plants. Soluble protein in-creased 43.48% at 400 Gy and 69.57% at 500 Gy g-ray ex-posures

Alikamanoglu et al.(2010)

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resulted in an increase in free fatty acids (FFA) levels. Adose-dependent decrease in the triacylglycerol content andconcomitant increase in free fatty acids was observed afterg-irradiation of nutmeg (Niyas et al. 2003). Similarly a pro-longed irradiation of seeds with UV light led to an increasein level of lipid peroxidation in wheat sprouts (Rogozhin etal. 2000). This suggested a breakdown of acylglycerol duringradiation processing, resulting in the release of free fattyacids (Niyas et al. 2003). Phospholipids, glycolipids, andneutral lipids were considerably reduced after g-irradiation,with concomitant reduction in the process of sprout growthin garlic sprouts (Pérez et al. 2007).

5. Oxidative stress and anti-oxidant defensesystemPlants often face the challenge of several environmental

conditions that include such stressors as drought, salinity, pes-ticide, low temperature, and irradiation, all of which exert ad-verse effects on plant growth and development (Foyer et al.1994). g-irradiation is reported to induce oxidative stress withoverproduction of reactive oxygen species (ROS) such assuperoxide radicals (O2

–), hydroxyl radicals (OH–) and H2O2(Apel and Hirt 2004), which react rapidly with almost allstructural and functional organic molecules including proteins,lipids, nucleic acids causing disturbance of cellular metabo-lism (Salter and Hewitt 1992). Hydroxyl radicals are gener-ated by ionizing radiation either directly by oxidation ofwater, or indirectly by the formation of secondary, partiallygenerated ROS. The generation of ROS is widespread in bio-logical materials and oxygen derived radicals include speciessuch as peroxyl (ROO·) and alkoxyl (RO·) radicals. One resultof oxidative stress is cellular damage by hydroxyl radical at-tack. The amount and rate of hydroxyl radicals generationfrom ROS is controlled partly by the cellular antioxidant sta-tus and partly by the availability of systems capable of reduc-ing (or ‘activating’) superoxide or hydrogen peroxide. It hasbeen shown that the effects of H2O2 resemble those of ioniz-ing radiation (Riley 1994). A major, direct target of g-irradia-tion that is probably the most important one is the watermolecule, which is omnipresent in organisms. The primary re-actions are excitation and ionization, which produce ionizedwater molecules (H2O˙+) and the radicals H˙ and ˙OH. How-ever, in biological tissue these ionizations are induced allalong the path of the radiation and lead to chain reactions,which produce secondary reactive oxygen species (ROS) as aresult of H˙ and eaq– becoming trapped. The most importantROS is H2O2; O2˙– is produced to very low extent, dependingof the O2 concentration (Tubiana 2008; Lee et al. 2009). The˙OH radical can react rapidly with various types of macromo-lecules, including lipids, proteins and, in particular, DNA.However, some of the resulting injuries can be readily re-paired and recovered, depending on the dose range. Differentworkers have described effect of g-irradiation on expressionof anti-oxidant enzymes as represented in (Table 6).

5.1 Superoxide dismutase (SOD; EC 1.15.1.1)Superoxide dismutase (SOD) isozymes are compartmental-

ized in higher plants and play a major role in combating oxy-gen radical mediated toxicity. In this review, we evaluate themode of action and effects of the SOD isoforms with respectT

able

5(concluded).

Dosageappliedandduratio

nof

irradiation

Species

Plantpartor

age/stage

ofplantg

rowth

forg-irradiatio

nPh

yiological

andbiochemical

response

References

100–

400Gy;

dose

rate

of0.864kG

y/h

Wheat

(Triticum

aestivum

L.)

Seeds

Totalchlorophyllincreased64.5%

at100Gy.

Prolinecontent

decreasedto

30%

Borzoueiet

al.(2010)

200,

700,

1200

Gy

Barley(H

ordeum

vulgare

spp.)

Seeds

50%

reductionin

shootlength

with

1200

Gy,

29.67%

de-

crease

inroot

numberwith

1200

Gy.

Nasab

etal.(2010)

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Table 6. Effects of the exposure to g radiation on antioxidant enzymatic and non-enzymatic components in different vegetal systems.

Dosage applied and durationof irradiation Species

Plant part or age/stage ofplant growth for g-irradiation Antioxidant defense response References

30 or 50 Gy (Nicotiana tabacum cv.Xanthi)

Young tobacco plants 6- to8-leaf stage

GST, sodCp (CuZnSOD), POD, Ngcat1 (CAT) expression in-duced at 30Gy.

Cho et al. (2000)

2, 4, 8, or 16 Gy; dose rate150 TBq

Red pepper (Capsicumannuum L. cv.Taeyang)

Seeds SOD activity was 2% to 25% and APX activity increaseddose-dependently from 27% to51%higher in the irradiationgroups than in the control, while that of GR was 56% to61% lower in the former, ascorbate content also increased ina similar manner, by 4% to 22%

Kim et al. (2005)

1, 2, 4, 6, 8, and 10 kGy;dose rate 228Gy/min

Soybean seeds (Glycinemax (L.) Merr.)

Seeds Decrease of CAT activity significant only at 100 and 120 Gyand the maximum decrease was detected at 120 Gy(62.4%), 140 Gy which caused a decrease in GSH-Px activ-ity by 33.7%, The GSH quantity increased as to 140 Gy(27.0% greater than the control) and then decreased at thehighest irradiation dose (36.0% less than the control), LPincreased by 21.6% and HO˙ quantity by 79.3% at 140 Gy.

Štajner et al. (2007)

2, 4, 8, 10, 12, and 16 kGy;dose rate 0.96 kGy/h

Kalungi (Nigella sativaL)

Seeds g irradiation enhanced the scavenging activity in acetone andmethanol extracts by 10.6% and 5.4%, respectively, at 16kGy.

Khattak et al. (2008)

0, 40, 60, 80, 100 Krad;dose rate of 233.5 rad/ min

Trigonella stellata, Tri-gonella hamosa andTrigonella anguina

Seeds CAT activity in shoots of T. hamosa decreased by 7%, 14%,34%, and 48% at 40, 60, 80 and 100 Krad, respectively ascompared with control. Induction of APOX and GR activ-ities in T. stellata roots was significant (P < 0.05) at 40Krad and high significant (P < 0.01) at 60, 80 and 100Krads

Al-Rumaih and Al Rumaih(2008)

2, 5, 10, 15, 20, 30, 50, 80,and 100 Gy; dose rate of0.54 Gy/min

Broad beans (Viciafaba L.)

Seeds g-rays of 20 Gy increased the activities of GR by 87.5%,SOD by 12.6%, APOX by 22.9%, and G6PDH by 38.9%and decreased H2O2 content by 17.8% as compared with thenonirradiated plants

Moussa (2009)

0.5, 2.0, and 5.0 kGy; doserate 0.09072kGy/min

Soybean seeds (Glycinemax (L.) Merr.)

Seeds Maximum relative enhancement of 80% in FRSA and 33% inFRAP at 2.0 kGy

Dixit et al. (2010)

2.5, 5.0, 10.0, and 20.0 kGy;dose rate, 6.5 kGy/h

Broad beans (Viciafaba L.)

Seeds H2O2 content and concentration of MDA significantly in-creased reached its maximum (36.3 and 38.2 mmol/g dw)compared to the control (2.3 and3.9 mmol/g dw) at dose le-vel 20.0 kGy

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to radiation induced oxidative stress resistance, correlatingage, species, and specificity of plants during development.Cells exhibiting high levels of SOD, catalase, and peroxidaseactivity are relatively less vulnerable to secondary effects ofradiation. Since superoxide is sufficiently stable to permit dif-fusion within the cell it is possible that it acts as an electrondonor to transition metals in irradiated tissue and may accountfor sensitization of cells to the effects of H2O2 by radiationexposure and the protection afforded by SOD to irradiationcells (Petkau 1987). Slooten et al. (1995) reported that thestimulation in SOD activity in response to stresses is possiblyattributed to the de-novo synthesis of the enzymatic protein.Changes in Peroxidase (POD) and superoxide dismutase iso-zyme compositions were seen in Vigna radiata (L.) R. Wilc-zek, as well as in calli of two tobacco species (Nicotianatabaccum and Nicotiana debneyi) after the irradiation (20–200 Gy) established from leaf discs (Roy et al. 2006; Wadaet al. 1998). Pramanik (1997) correlated morphological dam-age caused by g-rays with the changes in SOD isozyme pat-terns in callus tissue obtained from irradiated seeds. SODactivity PAGE gels showed an extra band (Rf value – 0.59)in all of the irradiated samples of calli, which were absent inthe control. This over expression of SOD is an indication ofradioprotection in vitro. Alteration in levels of SOD isoformshave also been correlated with effects of UV-B radiation,ozone, paraquat, methyl viologen (Krupa and Kickert 1989;Runeckles and Krupa 1994). Over expression of pea chloro-plast Cu/Zn SOD in tobacco leaves can improve their photo-synthetic performance under moderate stress and maintainphotosynthetic capacity even after severe oxidative stress(Gupta et al. 1993). The stress induced by g-irradiation mighthave imparted a shift in gene expression, which resulted information of new bands. A similar observation has been re-ported by El-Farash et al. (1993) who showed that inductionof such protein bands depends upon intensity of the stressand is genetically controlled. The stimulation of SOD activityis possibly due to a positive regulation of SOD genes or ofone particular encoding allele, in response to g-irradiationstress as shown in different biological models (Inzé and VanMontagu 2002). However, the disappearance of some bandsin irradiated samples immediately after exposure to g-rayscould be due to either degradation (Sen Raychaudhuri andDeng 2000) or to the switching off of the transcription–trans-lation machinery during radiation exposure and it gettingturned on again in the irradiation recovery phase. It has beenpropounded by (Zaka et al. 2002) that change in antioxidantenzymes must be directly linked to protein synthesis. Ascor-bate peroxidase (APX) and SOD activities in peppers also in-creased at low doses (from 2 to 8 Gy), whereas glutathionereductase (GR) activity decreased (Kim et al. 2004, 2005).Catalases (CAT) were stimulated at 5.16 C kg–1 Singh (1974)in safflowers. SOD, POD, CAT, and G6PDH (glucose-6P de-hydrogenase) activities are strongly stimulated in the case ofnew chronic external irradiation. Thus, it has been suggestedthat these enzymes protect plants more efficiently than the as-corbate glutathione ROS detoxification cycle.

5.2 Peroxidase (APX;EC 1.11.1.11) and Catalases (CAT;EC 1.11.1.6)Plant exposure to g-irradiation brings about changes in bi-

ochemical activity through various metabolites produced as

function of g-irradiation. The most important of these metab-olites are peroxyl radicals. The accumulation of organic per-oxides and oxidation of membrane lipids place a stress oncellular activity (Mead 1976). In plants, physiological effectsof g-irradiation were recognised by Ikeya et al. (1989); Voi-sine et al. (1991) to have caused the production of free per-oxyl radicals through the peroxidation of unsaturated fattyacids. The general equation for peroxidases activity is:H2O2 + DH2→ 2H2O + DHydrogen peroxide deposits were clearly increased in

pumpkin leaves and petioles at the high acute dose of 1 kGy(Wi et al. 2006a, 2006b). These deposits were present in thevessels, in the plasma membrane, in the cell corners of themiddle lamella and especially in the parenchyma cells. Thisaccumulation is concomitant with an increase in POD activ-ity in the corners of the middle lamella, where H2O2 is elim-inated (Wi et al. 2007). This stimulation of POD activity wasalso observed in garlic bulbs at low doses (10 Gy) (Croci etal. 1991, 1994), in Saintpaulia petioles at high-doses (0.85and 1.98 C kg–1;Warfield et al. 1975) and in sweet potatoroot disks (900 Gy;Ogawa and Uritani 1970). The radiationprotective effect of peroxidase is due to removal of H2O2,peroxides, and especially lipid hydrogen peroxides. This ac-counts for the greater effectiveness of peroxidases than cata-lase (Croute et al. 1982). Khanna and Maherchandani (1981)observed stimulated growth and increase in peroxidase activ-ity at low doses of g-irradiation in chickpea. g-irradiation caninfluence the isozymatic composition of peroxidases, as dem-onstrated by various authors in other species (Endo 1967;Ogawa and Uritani 1970; Shen et al. 2010). The study of per-oxidase activity will contribute to an understanding of mech-anism involved in radiation induced inhibition of plantgrowth. The peroxidase isozyme pattern was related to dam-age caused by irradiation (Shen et al. 1991). g-irradiation ofcallus culture of Datura innoxia Mill. resulted in stimulationof peroxidase activity and particularly increased the peroxi-dase enzymes with high electrophoretic mobilities (Jain et al.1990). The activity and isozyme of POD in Nicotiana deb-neyi Domin. and Nicotiana tabacum L., SOD in Nicotianadebneyi Domin., and CAT in Nicotiana tabacum L. increasedin response to g-irradiation treatment (Wada et al. 1998). La-ser irradiation of Vicia faba L. seeds could raise SOD, POD,and CAT activities and could eliminate the accumulation ofpoisonous free radicals and prevent lipid peroxidation (Alyand El-Beltagi 2010). In N. tabacum L., the genes GST, Cu/Zn SOD, POD, and CAT are up-regulated and the gene forcytosolic and stromal APX is down-regulated (Cho et al.2000). Indeed APX, GR, and MDHAR (monodehydroascor-bate reductase) activities were found to be either poorlystimulated or not stimulated at all (Calucci et al. 2003).ROOH catalase → H2O + ROH + A2H2O → 2H2O + O2Chaomei and Yanlin (1993) reported an increase in the ac-

tivity of POD and CAT with a corresponding decline ingrowth of Triticum aestivum L. plants under higher irradia-tion doses (20, 40, 60, 80 Kr). Singh et al. (1993) reportedinduction of APX activity in two sugarcane varieties grownunder g-irradiation. The activities of POD, CAT, and SODin radish (Raphanus sativus L.) leaves were enhanced by g-irradiation (10 Gy) treatment. Inhibition of CAT activity wasalso reported under irradiation stress (Ye et al. 2000; Štajner

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et al. 2009; Vandenhove et al. 2009). The present increase inAPX activity was reported to compensate for the progressivedrop in catalase activity. Peroxidase was considered to be thekey enzyme for the decomposition of H2O2, especially underCAT inactivation. Pasternak (1987) attributed peroxidase ac-tivation to membrane injury and the resulting shift in cellularCa2+ levels. According to Karpinski et al. (1997) APX acti-vation in Arabidopsis thaliana (L.) Heyn subjected to oxida-tive stress occurred through induction of APX1 and APX2gene transcription. Zaka et al. (2002) further reported en-hanced expression of APX gene in cells undergoing lowchronic g-irradiation stress. Several studies attributed enzymeinduction either to up-regulation of encoding genes or to ac-tivation of existing enzyme pools (Foyer et al. 1997) by amodulatory effect of enzyme structure.

5.3 Glutathione Reductase activity (GR; EC 1.6.4.2)GR activity in shoots and roots of the three Trigonella L.

genus significantly increased above the control values whenexposed to different doses of g-rays. Increased activity ofthis enzyme has been reported earlier in cotton when sub-jected to elevated atmospheric O2 (Foster and Hess 1980), inMg2+ deficient bean leaves, and in peas fumigated withozone (Hurkman and Tanaka 1987). Higher GR activity ofsalt-stressed cotton was reported to be due to an increase inglutathione turnover rate (Gossett et al. 1991). Foyer et al.(1995) reported an increase in GR activity in higher plantsas a result of enhancement of the transcription rate of encod-ing genes. GR would also be involved in the recycling of theelectron donor (glutathione, GSH) later in the pathway. Therewas slight increase in GR activity in Stipa capillata L. whenexposed to g-irradiation, probably due to enhancement of thetranscription rate of encoding genes (Foyer et al. 1991). Thehypothesis of GR radio-induction in Stipa is supported by thefact that a similar response was reported by other workers(Zaka et al. 2002; Kim et al. 2004; Moussa 2006). The roleof GR in H2O2 scavenging mechanism in plant cells was wellestablished in Halliwell–Asada enzyme pathways (Yamane etal. 2009).

5.4 Malondialdehyde (MDA) content and llipidperoxidationLipid peroxidation refers to the process whereby free radi-

cals “steal” electrons from the lipids in cell membranes, re-sulting in cell damage. This process proceeds by a freeradical chain-reaction mechanism. Thiobarbituric acid reac-tive substances (TBARS), the cytotoxic product of lipid per-oxidation, is normally considered as the major TBA-reactingcompounds that indicate the magnitude of the oxidative stress(Qadir et al. 2004; Qureshi et al. 2007). The basic effect ofradiation on cellular membranes is believed to induce lipidperoxidation by the production of free radicals (Leyko andBartosz 1986). Lipid peroxidation products in leaves of Ara-bidopsis thaliana L. present highest at full flowering and de-creased with higher g-exposure at this growth stage. At theother two growth stages, lipid peroxidation products were un-affected by g-treatment (Vandenhove et al. 2009). The Ma-londialdehyde (MDA) content and HO˙quantities wereobserved only under the highest irradiation dose, in soybean(Glycine max Merill.) seeds. The MDA quantity increase of

21.6%, and HO˙ radical quantity increase of 79.3% comparedwith the non-irradiated control (Štajner et al. 2009).

5.5 AscorbateAntioxidants are regarded as compounds that are able to

delay, retard, or prevent oxidation processes. They can inter-fere with oxidation by reacting with free radicals, chelatingmetals, and also by acting as oxygen scavengers, triplet aswell as singlet form, and transferring hydrogen atoms to thefree radical structure (Kitazuru et al. 2004). Ascorbate hasbeen shown to have an essential role in several physiologicalprocesses in plants, including growth, differentiation, and me-tabolism (Foyer 1993). The effect of ionizing radiation on vi-tamin C content has produced contrasting results for bothstrawberries and potatoes. In the case of strawberries, eitherno effect (Beyers et al. 1979) or a decrease in ascorbic acidcontent (Graham and Stevenson 1997) has been reported.Although some loss of vitamin C has been observed in pota-toes soon after irradiation (Maltseva et al. 1967), there wereno significant differences between irradiated and non-irradi-ated samples after a period of prolonged storage (Salkova1957). On the other hand, several studies have reported a re-duction in ascorbic acid content of potatoes following irradi-ation and storage (Joshi et al. 1990). However, it has beennoted that when reporting vitamin C levels, a number ofworkers have not taken into consideration the fact that ioniz-ing radiation can cause partial conversion of ascorbic acid orascorbate to dehydroascorbic acid (DHAA) (Diehl 1990; Kil-cast 1994). Biosynthesis of ascorbic acid and riboflavin inradiated corn, chickpea and soybean was found to more thanin control samples during germination. Low doses of g-irra-diation in addition to growth catalyzing enzymes or hor-mones would have accelerated the metabolic activity ofascorbate during germination just like X-rays (Sattar et al.1992).

5.6 Glutathione (GSH)Upon the imposition of oxidative stress, the existing pool

of reduced glutathione is converted to oxidized glutathione(GSSG) and glutathione biosynthesis is stimulated (May andLeaver 1993; Madamanchi et al. 1994). Together with ascor-bate, the most abundant antioxidant in plants, glutathionecontributes to protect plant tissues by direct scavenging of ac-tivated oxygen species (Halliwell and Gutteridge 1989).DHARDehydro-ascorbate + 2GS → Ascorbate + GSSGGSSG + NADPH → 2GSH + NADPRecently, there has been considerable interest in mecha-

nisms for diminishing the GSH levels in cells as a radio sen-sitization strategy (Riley 1994). Corn seedlings responded tog-irradiation by increasing the level of the glutathione perox-idase activity and showing no dependence on the genomecomplexity (Marchenko et al. 1996). The decrease in ferricreducing ability of plasma (FRAP) value in soybean seeds in-dicated a decrease of non-enzymatic antioxidants Vitamin Cand E and polyphenol constituents (Benzie and Strain 1996).The GSH quantity increased in soybean seeds as irradiationdose increased to 140 Gy and then decreased at the highestirradiation dose, which is a hermetic type of response underapplied doses of irradiation (Štajner et al. 2009). The amountof reduced glutathione, which is one of the most important

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non-enzymatic antioxidants, was positively affected by g-irra-diation. It has been reported that low level doses of g-irradi-ation enhance glutathione reductase activity (Chakravarty andSen 2001). However, no effect of irradiation was observed onconcentration or reduction state of the nonenzymatic antioxi-dants, ascorbate and glutathione in Arabidopsis thaliana L.(Vandenhove et al. 2009).

6. Effect of γ-radiation on bioactive agentsand oil componentsMany of the plant-derived phenolic compounds (flavo-

noids, isoflavonoids, coumarins, and lignans) are secondaryproducts of phenylpropaniods metabolism (Dixon and Paiva1995; Douglas 1996). In higher plants, phenylpropaniodsmainly hydroxycinnamic acid, cinnamoyl esters, flavones,flavonols, and anthocyanins provide a UV-A and UV-Bscreen. Both soluble and insoluble phenylpropaniods absorbefficiently in the range of 304–350 and 352–385 nm, respec-tively. Concerning the effect of g-irradiation on contents andcomponents of volatile oils, different results were reported.The differences were mainly due to the species of the plantand the dose applied. Koseki et al. (2002) studied the effectof radiation doses (0, 10, 20, and 30 kGy) on the flavonoids,essential oils, and phenolic compounds of Brazil medicinalherbs. From the described pharmacological tests of Brazilianmedicinal herbs carried out during this study, it was con-cluded that phytotherapy showed identical therapeutic actionas non-irradiated preparations after exposure to a dose of 10,20, and 30 kGy of ionizing radiation. The irradiation of tradi-tional medicines and herbal products does not result in anynegative chemical changes or important losses of active com-ponents. After irradiation up to 17.8 kGy, the content of themain biologically active substances of two medicinal herbs(ginkgo and guarana) were not modified (Soriani et al.2005). Mishra et al. (2004) reported that the dose of 5 kGyled to a decrease in 6-gingerol, the compound responsiblefor the pungency of ginger. Lee et al. (2005) showed that thepungency and red colour caused by capsanoids and capsan-thin, were not altered when irradiated (3, 7 and 10 kGy) inred pepper powder. Effect of g-irradiation at 10 kGy on thefree radical and antioxidant contents in nine aromatic herbsand spices (basil, bird pepper, black pepper, cinnamon, nut-meg, oregano, parsley, rosemary, and sage) were studied by(Calucci et al. 2003). Irradiation resulted in a general in-crease of quinone radical content in all of the investigatedsamples, as revealed by Electron paramagnetic resonance(EPR) spectroscopy, and in a significant decrease of total as-corbate and carotenoids content of some herbs. Polovka et al.(2006) reported that irradiation (5 to 30 KGy) of groundblack pepper shows some significant influences on the anti-oxidant activities with respect to the non-irradiated samples.The most significant changes of antioxidant activity were ob-served in creation of thiobarbituric acid reactive substances.Irradiation with 15 kGy caused slight increase in phenol con-tent in Brassica nigra L. Koch (0.1%) followed by Cassiasenna L. (pods) (1.3%). However, the maximum increasewas about 70% and was observed in Cassia senna L. (leaves)followed by Lepidium sativum L. (25.6%) and Cymbopogonschoenanthus L. (24.9%). On the other hand irradiation with15 kGy reduced the phenol content of Trigonella foenum-

graecum L. by 4.1% followed by Hibiscus sabdariffa L.(5.1%) and Acacia nilotica L. (14%). The maximum reduc-tion was 33% and was observed in Cymbopogon citrates L.The effect of g-irradiation on the phenolic content of theplants under study has not been investigated previously, butsimilar observations on other biological materials were re-ported. Adamo et al. (2004) reported an increase in phenolcontent for irradiated samples of truffles at the dose level inthe 1.0–1.5 kGy and proposed that the destructive process ofoxidation and g-irradiation were capable of breaking thechemical bonds of polyphenols, thereby releasing solublephenols of low molecular weight. In a study on nutmeg, Var-iyar et al. (1998) found that the essential oil constituentsshowed clear quantitative differences upon g-irradiation.Thus, the content of á-terpeniol, 1-terpinene-4-ol, and myris-ticin was increased in irradiated nutmeg, while that of sabi-nene, a-pinene, and elimicin decreased in comparison to thecontrol sample. Less significant changes in the content ofsome of the essential oil constituents between the controland the irradiated samples were noted in clove and carda-mom. Seo et al. (2007) showed that among the essential oilsconstituents of Angelica gigas Nakai, oxygenated terpenessuch as a-udesmol, a-eudesmol, and verbenone were in-creased after irradiation, but their proportions were variablein a dose dependent manner.Breitfellner et al. (2002) studied the effect of g-irradiation

on flavonoids in strawberries and reported that in hydrolyzedsamples four phenolic acids (gallic acid, 4-hydroxybenzoicacid, p-coumaric acid, and caffeic acid) were identified andfive flavonoids were detected in hydrolyzed samples ((+)-cat-echin,(-)-epicatechin, kaempferol-3-glucoside, quercetin-3-glucoside, and quercetin-3-galactoside). The concentration of4-hydroxybenzoic acid increased and that of catechins andkaempferol-3-glucoside decreased as irradiation dose in-creased, whereas those of quercetin-3-glucoside remained un-changed up to a dose of 6 kGy. Accumulation of phenoliccompounds in cells is demonstrated and may be explainedby the enhancement of phenylalanine ammonia lyase (PAL)activity. Bhat et al. (2007) reported that some compoundssuch as phytic acid in velvet bean seeds (Mucuna pruriensL.) was completely eliminated on exposure to a dose of 15kGy. Decrease or elimination of phytic acid is likely due tothe degradation of phytate to lower inositol phosphates andinositol by the action of free radicals generated during irradi-ation. Another possible mode of phytate loss could have beenthrough cleavage of the phytate ring itself (Duodu et al.1999). Similarly, treatment of broad bean (Vicia faba L.)seeds with g-irradiation reduced the level of phytic acid com-pared to controls (Al-Kaisey et al. 2003). Significant quanti-tative changes were noted by Variyar et al. (1998) in some ofthe phenolic acids in clove (Syzygium aromaticum L.) andnutmeg upon irradiation with 10 kGy dose. They reportedthat the content of gallic and syringic acids in irradiatedclove was increased, whereas that of p-coumaric, ferulic, andsynapic acids decreased to approximately half in the controlspice, and that of caffeic and gentisic acids remained un-changed. In the case of irradiated nutmeg (Myristica fragransL.), except for protocatechuic acid and p-coumaric acid,which remained unchanged, the content of other phenolicacids showed wide variations compared with that of controlsamples. Elevation of some compounds by g-irradiation may

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Table 7. Effects on g irradiation on the oil and metabolite content of different species.

Dosage applied and duration ofirradiation Species

Plant part or age /stage ofplant growth for g-irradiation Metabolites and oil content/yield References

5, 10, and 15 kGy; dose rate of0.98 kGy/h

Rice (Oryza sativa L.) Bran The loss of total E vitamers and oryzanol occurred intwo stages: 50%–82% and 12%–33% immediatelyfollowing 15kGy irradiation.

Shin and Godber (1996)

1, 2, and 5kGy Mushroom (Agaricus bisporus) Fruiting body Eight-carbon compounds decreased as the doses of g-irradiation increased, from 41.73 for the control (0kGy) to 20.06 (1 kGy), 8.77 (2 kGy), and 4.04 mg/g (5 kGy irradiated mushrooms)

Mau and Hwang (1997)

0.3 kGy; dose rate 6kGy/h Citrus fruit (Citrus clementiaHort.ex. Tanaka)

Seed Increment and accumulation of scoparone and PALactivity increased to 250 pKat/gdw

Oufedjikh et al. (2000)

0.89, 2.24, 4.23, and 8.71gGy;dose rate 0.1 kGy/min

Orange (Citrus limonia) Orange juice Myrcene, linalool, geranial and neral decreased atrate of 5.86%, 5.66%, 5.83%, and 6.54% per kGydose applied. At 5-D dose myrcene, linalool, neraland geranial would be reduced by 20.9%, 20.1%,20.7%, and 23.2%.

Fan and Gates (2001)

0.5, 1, 2, and 5 kGy; dose rate14Gy/min

Soybean (Glycine max Merrill) Seeds Decrease in diazdin (33.95), glycitin (37.14) and gen-istin (28.34)%, total glycone reduced to 26.97% at 5kGy.

Variyar et al. (2004)

2, 16, and 32 Gy Lithospermum erythrorhizon Seed, seedlings The g-irradiation significantly stimulated the shikoninbiosynthesis of the cells and increased the total shi-konin yields (intracellular+extracellular shikoninyields) by 400% at 16 Gy and by only 240% and180% at 2 and 32 Gy, respectively

Chung et al. (2006)

10, 20, 40, 60, 80, and 100kGy

Maytenus aquifolium Martius Leaves 5.2% decrease in tannic acid at 100 Gy. Campos et al. (2005)

1, 3, 5, 10, and 20 kGy Welsh onion (Allium fistulosumL.)

Bulbs Enhancement in total concentration of volatile com-pounds by 31.60% and 24.85% at 10 and 20 kGy,respectively.

Gyawali et al. (2006)

5 and 10 kGy Dry shiitake (Lentinus edodesSing)

Seeds The ratio of the eight carbon compounds, such as 3-octanone, 3-octanol and 1-octen-3-ol, to total vola-tiles decreased from 72% in the control to 21% inthe 10 kGy irradiated samples.

Lai et al. (1994)

1, 3, 5, 10, and 20 kGy Angelica gigas Nakai Leaves The major volatile compounds were identified 2,4,6-trimethyl heptane, a-pinene, camphene, a-limo-nene, beudesmol, a-murrolene and sphatulenol.The irradiated samples at doses of 1, 3, 5, 10 and20 kGy were 84.98%, 83.70%, 83.94%, 82.84%,and 82.58%, respectively

Seo et al. (2007)

2, 4, 8, 10, 12, and 16 kGy,dose rate 0.96 kGy/h

Kalungi (Nigella sativa) Seeds Extraction yields increases were 3.7%, 4.2%, 5.6%and 9.0% for hexane, acetone, water and methanolextracts. Phenol content increase from 3.7 for con-trol to 3.8 mg/g for 16 kGy.

Khattak et al. (2008)

1, 3, 5, 10, and 20 kGy Licorice (Glycyrrhiza uralensisFischer)

Root 10 kGy dose irradiation induced the maximum levelof total yield of volatile compounds by 12% butdecresed slightly at 20 kGy

Gyawali et al. (2008)

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be attributed to their higher extractability. Some reports indi-cate that irradiation decreases tannins in bean seeds (Villavi-cencio et al. 2000). Such differences may be attributed to thedifferential response and variability in the genetic constitu-ents (strains and varieties). Doses of 7.0 and 10 kGy signifi-cantly reduced the tannin content of Shahalla sorghumvariety but not that of Hemaira variety (Siddhuraju et al.2002). Various studies on effects of g-irradiation on activeprinciples in plants is summarized in Table 7.

7. Perspectives(g) radiations can be used to enhance the rate of genetic

variability because the spontaneous mutation rate is veryslow,which previously has prevented breeders from usingthem in plant breeding programs. Advanced molecular toolsused along with traditional plant breeding can improve cropproduction. Efforts in the future should be directed at meta-bolically engineering plants for the production of secondarymetabolites that confer resistance thereby providing an oppor-tunity for stable genetic character to confer radio resistance.Elicitation of secondary metabolites has applications in overproduction of desired compounds, which is an area of com-mercial importance especially for high-value low-volumeproducts. In this category belongs the production of antitu-mor and anticancer compounds that are of great medicinalvalue. As the plant is exposed to a number of external stim-uli, studies on signal transduction in plants for various phys-iological responses would require focused attention tounderstand its adaptation. g-rays can be used for inducingnew mutants using local germplasm enhancement for devel-oping new mutant varieties as well as for basic research ingene discovery and functional genomics. It is difficult to pre-dict the impact of climate changes on global, regional, or na-tional agriculture and therefore new varieties must bedeveloped and distributed regularly at the national and re-gional levels for sustainable crop production. g-rays can beemployed to develop new varieties that can be readilyadapted in a short period in different locations with varyingagro-climatic and growing conditions, and where availableresources are limited.

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2008. Effect of gamma radiation and gibberellic acid ongermination and alkaloid production in Atropa belladonna L.Aust. J. Basic Appl. Sci. 2(3): 401–405.

Abdel-Hady, M.S., and Ali, Z.A. 2006. Effect of gamma irradiationon wheat immature culture regenerated plants. J. Appl. Sci. Res.2(6): 310–316.

Abdullah, T.L., Endan, J., and Nazir, B.M. 2009. Changes in flowerdevelopment, chlorophyll mutation and alteration in plantmorphology of Curcuma alismatifolia by gamma irradiation.Am. J. Appl. Sci. 6(7): 1436–1439. doi:10.3844/ajassp.2009.1436.1439.

Al-Jassir, M.S. 1992. Chemical composition and microflora of blackcumin (Nigella sativa L.) seeds growing in Saudi Arabia. FoodChem. 45(4): 239–242. doi:10.1016/0308-8146(92)90153-S.

Al-Kaisey, M.T., Alwan, A.K.H., Mohammad, M.H., and Saeed, A.H. 2003. Effect of gamma irradiation on antinutritional factors inbroad bean. Radiat. Phys. Chem. 67(3–4): 493–496. doi:10.1016/S0969-806X(03)00091-4.T

able

7(concluded).

Dosageappliedanddu

ratio

nof

irradiation

Species

Plantpartor

age/stage

ofplantg

rowth

forg-irradiatio

nMetabolitesandoilcontent/y

ield

References

50,8

0,110,

and150Gy;

dose

rate

0.54

Gy/min

Atropabella

donnaL.

Seeds

Seedsirradiated

at110Gypossessed4.01

mg/gand

109.67

mg/planttwicealkaloid

valuethan

control

(2.03mg/gand53.41mg/plant).

Abdel-H

adyet

al.(2008)

2,4,

8,10,1

2,and16

kGy;

dose

rate

0.96

kGy/h

Kalungi

(Nigella

sativa)

Seeds

Extractionyields

increaseswere3.7%

,4.2%,5

.6%,

and9.0%

forhexane,acetone,

water

andmethanol

extracts.Ph

enol

contentincrease

from

3.7forcon-

trol

to3.8mg/gfor16

kGy.

Khattaket

al.(2008)

0,10

,20,

30,40,

50,6

0,and70

Gy;

dose

rate

4.64

kGy/h

Orthosiph

onstam

ineus

Shoottip

sRosam

arinic

acid

contentwas

lowest5.27

mg/gfw

inplantletsirradiated

at10kG

yandhighest8.40

mg/g

fwat

30Gy.

Kiong

etal.(2008)

5,10,a

nd15

kGy,

The

activ

ityof

thesource

was

6.345K

ciandtheenergy

1.25

MeV

Cam

elhay(Cym

bopogonschoe-

nanthus)

Leaves

21%

reductionin

tanninsas

aresultof

girradiation

andphenol

contentby

morethan

25%

at15

kGy

Musaet

al.(2010)

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