2 Review of

Literature
Rice is a member of grass family. It belongs to genus Oryza of

Gramineae family. The genus posses about 24 species out of which

22 are wild and two species namely Oryza sativa and Oryza

glaberrima are cultivated. While Oryza glaberrima was grown in the

tropical and subtropical Asia but center of domestication is a matter

of discussion. Some believes that simultaneous domestication in

various center extending from the plains below eastern hills of

Himalayas. Another view postulates a more limited center of

origin. Notably inland valleys in Thailand, Burma and Laos ,Oryza

glaberrima is generally considered as a wild specie .It is commonly

found in West Africa in a few secondary centers. As far as production

is concerned China is the top producer with 166,000.000 metric

tonnes, followed by India with 133,513,000metric tonnes. Brazil

posses lowest of production with 10,219,300 metric tonnes

( Anonymous,2009)

From the initial point or points of cultivation, Oryza sativa has

spread over a considerable part of the earth and has become a major

food crop for a larger part of world’s population. Varieties adaptation

generally includes development of the main subspecies i.e “Indica”

and “Japonica” and the improvement of land and water management

practices, which changed and optimizes rice edaphic and climatic

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environment. Rice is essentially a crop of sub-tropic and on the

higher elevations up to 6000 feet above sea level. Rice need

abundant supply of fresh water for irrigation followed by high

temperatures and high atmospheric humidity .Rice thrives over a

wide range of soils ranging from 4.5 to 8.5 pH. Only a few rice

varieties posses tolerance to saline and alkaline condition.

Rice is a staple food for 17 countries and at present it is grown

over more then 100 countries. Archaeological evidences indicated

that the sophisticated rice cultivation system existed in China over

7,000 years ago. Rice is grown in all the continents except Antarctica.

It is grown in 15.294m millions ha . Today rice produces food for

nearly 2/3 of world population about 4/5 of rice is produced by small

scaled farmers and is an source of income for 100 millions

households of Asia and Africa

RICE IN INDIA :- In India, rice is an important staple food for more

than 2/3rd of population. Rice provides a vital role in our national food

security and is a mean of livelihood for millions of rural households. In

India, a number of festivals such as Bihu, Pongal, Onam are

associated with rice harvest .

There is a considerable increase in productivity of rice in India

during the recent past. The productivity of rice was 668 kg/ha in

1950-51 has reached to 2,066 kg/ha during 2000-02. During the

period of 1961 to 2005, the cultivation of rice has increased from 35

m ha to 44m ha followed by increase in production from 54 million

tonnes to 124 million tonnes and productivity from 1.54 tonnes to

Review of Literature
2.93 tonnes/ha. This increase is basically due to cultivation of the

hybrid rice varieties in major areas with advanced rice production

technologies and mechanization ( Fairhust and Doberman, 2002

b). In India rice is the major staple food for 65% of total population.

The rice cultivation area extends throughout India. Rice

cultivation can be taken up in areas lying below sea level (Kerala)

and up to altitude of 2000m (Kashmir) . During last 45 years (1967 –

2008), the rice area has increased by one half times from 115-50

million ha to about 153.26 million hectares. It is estimated that the

rice demand in 2025 will be 140 million tonnes in India. The demand

for food grain is expected to rise not only as a function of population

growth but also as more and more people crosses poverty line.

Rice ranks second in position in terms of area harvested and

cultivated. In terms of importance as a food crop or in terms of

calorific value rice crop occupies first place amongst cereal crops.

The nutritive composition varies with environmental condition, the

germ, pericarp of aleurone layer are rich in endosperm. Carbohydrate

of rice is starch constituting about 72% to 75% with fiber content of

hemicellulose of pentose, Arabinose and Xylose. Protein “Glutelin”, is

also known as “Oryzenin”, rice also contains albumin, globulin and

prolamines. Rice is however , deficient in Lysine and Threonine.

Rice: Improvement Via Tissue Culture.

Plant tissue culture has become thrust area in all areas of crop

improvement programmes. The beginning of the plant tissue culture

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was made early in 1898, when a German botanist, G. Haberlandt

successfully cultured fully differentiated individual plant cells,

isolated from the different tissues in several plant species. During

1934 to 1939, due to discovery of the important hormones like

auxins and B vitamins, the foundations of the plant tissue culture

were laid down by three scientists (Gautheret, White and

Nabecourt.).

In vitro culture of rice begin in the early 1950 s, it was

“Fujiwara” and “Ojima” (1954) who first of all reported successful

culture of rice roots excised from seedlings germinated aseptically.

However, a major realization in this course of practice is the genetic

variability. This variation in tissue culture raised plants is proving to

be an rich source of crop improvement. Among the techniques anther

culture, protoplast fusion, leaf culture, root culture, immature embryo

culture and seed culture are important in course of crop

improvement.

Rate of callus induction generally depends upon various factors

that includes culture environment genotype, media composition,type

of explant and partial dessication.

( Vasil and Hildebrandt, 1965)

For the rice, the seeds derived embryogenic calli are

considered as most appropriate source of genetic variation (De

Datta et al., 1990). Embryogenic calli are convenient and large

quantity can be made with absolute uniformity in physiological

characters. They are now widely used for experimental purposes.

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 The genotype plays a very crucial role in callusing ability of

seeds (Abe and Futsuhara,1986) that requires different types of

medium i.e MS (Murashige, and Skoog,1962 ), N6 ( Chu, 1975),

B6 with different composition of growth hormone. However, for the

rice 2,4-D is found to be only growth regulator for callusing . In

addition to this there are various other chemical like casein

hydrosylate for Japonica rice and Indica Rice. Proline addition in the

medium also induces the callusing in various varieties. Sucrose

concentration however, plays an important role in callusing from the

anther (Guha and Mukherjee., 1964).

Variations induced in Culture :- The variation observed in tissue

culture is due to physiological changes induced by the culture

condition. However, sometimes these variations are temporary and

disappear when culture conditions are removed. However, sometimes

it persists for longer period. The first reports of morphological

variation observed in plants were published by Heinz and Mee

(1971).Since then, several useful variants of sugarcane resistant to

salt stress fungal and viral diseases have now been released. This

variation so produced through tissue culture could be useful for the

development of new cultivars ( Larkin and Scowcroft., 1981)

Genetic variants selected through tissue couture are referred to

as calli clones, protoclones from protoplast cultures. Somaclonal

variations are used to describe genetic variation regenerated from

the cell cultures . Plants regenerated from cell culture of gametic

origin are termed as “Gametoclones”.

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 To be of agronomic use, a somaclone must fulfill the basic

requirements i.e It must involves useful characters, it should be

superior from the parent, must be combined with all other desirable

characters of parents and the variations must be inherited stably

through successive generations.

Underlying principle behind the somaclonal variations lies in

quantitative phenotypic variations, activation of transposable

elements and high frequency of sequence changes (Bayliss ,1980).

The “DNA methylation” have been suggested as a possible

mechanism (Kaepler and Phillips, 1993) and patterns of DNA

methylation have been charaterised for many varieties (Selker and

Stevens,1985). The DNA methylation leads to chromatin changes

that ultimately cause disturbed replication timing and base changes.

Essential Role of Phosphorus (P) in plant

Phosphorus is an essential nutrient both as a part of several

key plant structural compound and as a catalyst in the conversion of

numerous key biochemical processes. Phosphorus is a vital

component of DNA, the genetic “Memory unit” of all living things. It is

also a component of RNA, which reads genetic code. The structures

of both DNA and RNA are liked together by phosphorus bonds.

Phosphorus is highly mobile in plant and when deficiency

occurs it may be translocated from old plant tissue to young actively

growing areas. Thus, early vegetative responses to phosphorus are

often observed .As the plant matures, phosphorus is translocated into

Review of Literature
the fruiting areas of the plant where high energy compounds are

needed for the formation of the seeds and fruits.

The percentage of total amount of the each nutrient taken up is

higher for phosphorus late in growing season than for either nitrogen

or potassium.The total phosphorus content of soils is generally very

low i.e 0.6% this compares to an average soil content of 0.14%

nitrogen and 0.83% potassium ( Hoford ,1997) , many factors

influence the content of soil phosphorus. Some of these are : (1)

Type of parent material from which soil is deserved ; (2) Degree of

weathering (3) Climatic conditions. In addition, soil phosphorus levels

are affected by erosion,crop removal and phosphorus fertilization

(Kirk et al., 1998).

Soil phosphorus is classified in two broad groups (1) Organic

phosphorus, found as plant residues, manure and microbial tissues.

In phosphorus rich soils 50% or more of the phosphorus is in organic

form compaired to 3% in deficient solis (2) Inorganic forms consists

of the apatite i.e original source of all phosphorus.

Soluble phoshorus either in the form of fertilizer or natural

weathering reacts with clay, iron and aluminium compound in the

soil and is converted readily to less available form by process called

“phosphorus fixation”.

Due to fixation, the phosphorus moves little in the soil (less

then an inch ) a crop absorbs more then 20% of fertilizer as a result

the phosphorus is lost by “Leaching”.

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 N P K Ca

Fig.1 Relative movement of nutrients in the soil

Precipitation of the phosphorus as slightly soluble calcium

phosphate occurs in calcareous soils with pH values around 8.0 under

the acidic condition, phosphorus is precipitated as Fe or Al

phosphates of low solubility, maximum availability generally occurs in

a pH range of 6.0 to 7.0 and this pH provide phosphorus as H2PO4-

that is readily absorbed by plant than HPO4-

(9.0) Tricalcium phosphates unavailable

7.0 Mono and dicalcium phosphates maximum available

( 5.5) Fe , Al phosphates unavailable

Fig.2 Effects of pH on phosphorus availability

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Phosphorus Deficiency : Effects on Crop-

The plant obtains phosphorus from the soil. It occurs in both

organic and inorganic forms. It is the most limiting element

after the nitrogen (Jungk , 2001) due to it's fixation in the

soil.

About 5-7 billion ha soil worldwide lacks the sufficient plant

available phosphorus (Bates, 1973) and almost 50% of rice soils are

currently phosphorus deficient. Phosphorus deficiency is widespread

in all major rice ecosystems and is major limiting factor in arid upland

soils where soil P fixation capacity is often very large.

( Raghothama, 1999).

The common causes of P deficiency are: low indigenous soil – P

supply, insufficient supply of mineral P fertilizers, low efficiency of

applied fertilizer, P immobilization ( Kirk et al, 1999), excessive

use of N fertilizers, crop establishment methods ( more likely in

direct- seeded rice where plant density is high and root system are

shallow).

Moderate P deficiency is generally different to recognize. In the

field, P deficiency is often associated with other nutrient disorders.

(Bagyarj and Verma, 1995).

Phosphors deficient plants are stumped with greatly reduced

tillering. leaves are narrow, short, very erect and dirty dark green.

Stems are thin and spindly and plant elongation is retarded. The

number of leaves, panicles and grain per panicle is also reduced red

Review of Literature
and purple colors formation occurs if variety posses property of

anthocynin production

(Doberman and Fairhost,2000b). Other effects include

delayed maturity, large production of empty grain, no response to

application of fertilizer.

Plant mechanism, adaptation for low phosphorus stress -

Plant have developed several mechanism to counter the

problem of low phosphorus availability, so as to acquire more

phosphorus from the surrounding rhizosphere. Such adaptations are

generally at the physiological biochemical and morphological levels.

Morphological adaptation for countering low P stress

General diffusion rates of phosphorus is very low because soil

particles easily binds the phosphorus .If P doesn’t move freely in the

rhizosphere, plant may increase P uptake by the expanding the root

system, thereby exploring more soil volume .The size of root system

is considered as an important factor of plant that tolerate P

deficiency ( Wissuwa,2002). Tolerance however, could be

achieved by the increase in the external P uptake efficiency, defined

as the phosphorus uptake per unit root size (RSA) (Raghothama.,

1999).

Alternatively, plant generally capable of releasing soil bound

phosphorus by excreting the organic compound, P solublization due

to organic ions ( anions) were responsible for the bulk of P uptake by

rice from P deficient soil ( Kirk et.al. 1999). Another response to low

Review of Literature
phosphorus availability is the symbiotic association with the

mycorrhiza. The fungus and plant show symbiosis where fungus helps

in aquitisation of phosphates (Bagyaraj and Verma,1995) . Fungus

so extends the phosphorus depletion zone (Harrison and Van

Burren ,1995. Generally genotypes with the more P uptake

efficiency are highly associated with higher relative root growth. This

is due to fact that additional P allows further biomass accumulation,

including root growth ,this is considered as primary factor for the

genotypic differences in P uptake ( Wissuwa, 2002).

It is also reported that under low phosphorus availability the

root to shoot ratio generally increases followed by increase in root

diameter with increase in absorptive surface area. The root

assimilates more root hairs as to enhance uptake of Pi. (Gahhonia

and Nielson. 1998).

Molecular and Biochemical mechanisms involved under

phosphorus limited environment :-

Efforts have been directed for improvement of tolerance to low

phosphorus stress. Since, there is considerable variation at genotypic

level for the ability to take up P from a highly fixing soil exists in gene

bank accession.

Genetic studies on tolerance to P deficiency in rice have

identified a major quantitaitve trait locus ( QTL) for Pi uptake

(Senanayake, 1984) which has subsequently been mapped to a

small marker interval on the long arm of chromosome 12.

( Wissuwa et al., 1998).

Review of Literature
 Two OsPHR genes from rice Oryza sativa .L were isolated and

designated as OSPHR-1 and OSPHR-2 based on the amino acid

sequences homology to AtPHR 1 ( Bari et.al., 2006) that plays an

crucial role during the Pi starved condition in Arabidopsis

(Hamberger et al ,2002). OsPHr – 1 and 2 are involved in Pi

starvation signaling pathway by regulation of the express of Pi-

starvation.

In E.coli there exists a high affinity phosphate transporter Pst

CAB – this is generally active transport system, more specifically on

ATP binding cassette ( ABC) transporter. It is generally activated

under the low phosphate levels. Similarly, OSP-11 have been

characterized (Wasaki et al., 2003 ). It is responsible for the

adaptation to early stages of adaptation to low phosphorus

environment ( Liu et al., 1997 ).

Recently OSPTF–1 (transcriptional factor) was introduced into

Nipponbare i.e a phosphorus deficiency sensitive, derived from

Kasalth (Keke et. al, 2005). The transformed cultivars showed an

(over expressing OSPTF–1) considerable tolerance to phosphorus

deficiency. Possibility of manipulation of the expression of

phosphorus transported gene may be involved in the mobilization of

phosphorus within soil.

In E.coli and Brewer's yeast there are reports of some groups

that are activated under phosphorus starved condition (Torriani

and Ludtke, 1990). Psi genes are useful in the enhanced uptake of

pi from soil under p starved condition. The PHO-regulon is now well

Review of Literature
known aggregation of gene ( Sacchromycese cerevisiae) that helps in

scavenging any available Pi from soil under starving condition ( Abel

et al, 2002).

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 The PHO regulon is generally regulated by the two component

system. namely PHO. B – Pho- R.

Pho R

(Autophosphorylation under P starved condition)

PhoB Enhanced uptake

Binds Pho–Box of Pho- regulon Activation of operon

Fig . Mechanism of Pho -regulon working (Torriani, 1990)

Similar to Pho ragulon of sacchromycese cerevisae, Pho.1 and

Pho. 2 and also Pho.3 have been isolated (Hamberger et al.,2002).

During the phosphorus starved state the common enzymes

associated with phosphate metabolism gets activated, (Green,

1994). This includes “Phytase ”, RNAase” and some phosphate

transporters (Raghothama, 1999). These enzymes plays

crucial role in phosphate nutrition during the stress by performing

phosphorylation and dephosphorylation thereby causing activation

and deactivation of many proteins specially transporters necessary

for phosphorus uptake ( Liu et. al., 1997).

Acid phosphatase is the most crucial enzyme that is involved in

the release of the Pi from organic compounds, ions, soils and tissues

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( Tabatabai and Bremner,1969). The activity of acid phosphatase

generally increase under the low phosphorus stress state. Acid

phosphatase is responsible for moblisation of Pi from phosphate

esters.

It is now well know that the low phosphorus stress leads to

accumulation of the polyamines specially “putrescine” ( Bertoldi et

al, 2004). It was found that common precursor of putrescin i.e.

Argenine decarboxylase complex (ADC) generally increases in various

stresses including P. This in turn increases synthesis of the putrescin

production. The decarboxylated form of SAM i.e S. adenosyl

methionine is precursor of the putrescin and ethylene. Thus, under

low P condition ethylene production is also elevated. Polyamines are

general indication to stresses like acid stresses and somatic stresses

( Flores and Galston, 1982).

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