transgenics for delayed fruit ripening

Transgenics For Delayed Fruit Ripening

By

Sukanya 1385

Highly coordinated

Genetically programmed

Irreversible phenomenon

Physiological biochemical changes

Development of a soft and edible ripe fruit

What is Fruit Ripening

The molecular mechanisms controlling the ripening of fruit

Open Archive TOULOUSE Archive Ouverte

Increased respiration

Chlorophyll degradation

Biosynthesis of carotenoids anthocyanins essential

oils Flavor and aroma Components

Increased activity of cell wall-degrading enzymes

Transient increase in ethylene production

What are the changes

Major Developmental Changes during Tomato Fruit

Development and Ripening

The Plant Cell Vol 16 S170ndashS180 2004

Based on their respiratory pattern and ethylene biosynthesis

during ripening

VPrasanna et al

Classification of fruits

Pathway for ethylene biosynthesis

Rate limiting step

Critical Reviews in Food Science and Nutrition 471ndash19 2007

The expression of

ethylene biosynthesis

and ethylene

perception genes

during the transition to

climacteric in tomato

Kevany et al 2007

Bleecker and Kende 2000

Ethylene Perception and Signal Transduction

Fruit pulp or the mesocarp

parenchymatus cells

complex network of polysaccharides and proteins

The primary cell wall contains

35 pectin

25 cellulose

20 hemicellulose

10 structural hydroxyproline-rich protein

Structural components of fruits

Plant polysaccharides play a major role in

storage mobilization of energy and in

maintaining cell and tissue integrity due to

their structural and water binding capacity

Enzymes Related to Pectin Dissolution

Critical Reviews in Food Science and Nutrition 471ndash19 (2007)

Globally cultivated fleshy fruit

Worldrsquos largest vegetable crop after potato

Indian production scenario-

350000 hectares 5300000 tonsyear

Short generation time 3-4 months

Simple genetics

Numerous characterized mutants

Cross fertile wild germplasm to promote genetic studies

Routine transformation technology

Postharvest losses-5 to 25 in developed countries

-20 to 50 in developing countries

Tomato model systems for fruit development

and ripening

Natural Mutants Affected in the Ripening Phenotype

Disadvantages of existing methods of storage

Labor intensive

costly

Occupies a large floor space

Poor heat transfer may occur resulting in poor product quality

Excessive dehydration in unpacked products

Chemical changes during freezing

-enzyme-activated browning

-development of rancid oxidative flavors

Textural changes during freezing

-mushy and watery

The use of 1-methylcyclopropene (1-MCP) on fruits and

vegetables

Inhibitor of ethylene perception

Easily released as a gas when the powder is dissolved in water

Approved by the Environmental Protection Agency (EPA) in 1999

Marketed as EthylBlocreg by Floralife Inc (Walterboro SC)AgroFreshInc a

subsidiary of Rohm and Haas (Springhouse PA)

CB WatkinsBiotechnology Advances 24 (2006) 389ndash409

Transgenic approach

Delayed fruit

ripening

BLOCKING THE PERCEPTION OF ETHYLENE

BLOCKING THE EXPRESSION

OF GENES INDUCED BY ETHYLENE

BLOCKING ETHYLENE SYNTHESIS

Regulation of Ethylene Production

a Suppression of ACC synthase gene expression

ACC (1-aminocyclopropane-1-carboxylic acid) (ACS2)

conversion of S-adenosylmethionine (SAM) to ACC

the second to the last step in ethylene biosynthesis

an antisense (ldquomirror-imagerdquo) or truncated copy of the synthase gene

Oeller et al 1991

Yao et al1999

Nath et al 2006

Antisense Technology

httpagbiosafetyunleduflashantisenseswf Journal of Plant Physiology170987ndash 9952013

Null Mutation of the MdACS3 Gene

Apple cultivars homozygous or heterozygous for null allelotype

showed no or very low expression of ripening-related genes and

maintained fruit firmness

Aide Wang 2009

RNAi-mediated silencing

Chimeric RNAi-ACS construct designed to target ACS

homologs

Delayed ripening and extended shelf life for sim45 days

Aarti Gupta Ram Krishna Pal Delayed ripening and improved fruit processing quality in tomato by RNAi-mediated silencing of three

homologs of 1-aminopropane-1-carboxylate synthase gene Journal of Plant Physiology 170 (2013) 987ndash 995


b Suppression of ACC oxidase gene expression

It catalyzes the oxidation of ACC to ethylene

The last step in the ethylene biosynthetic pathway

Down regulation through anti-sense technology

Hamilton et al 1990

Ye et al 1996

Xiong et al 2003

Ripening in papaya fruit is altered by ACC

oxidase cosuppression

Fig1Map of the construct pKYCPACOO-1 containing the ACC oxidase

fragment cloned in PKYLX80 in the sense orientation The ACC oxidase

fragment is flanked by the CaMV 35S promoter and the RUBISCO

terminator

Fig2 Ethylene production in papaya transgenic fruits

Rodolfo Loacutepez-Goacutemezet alTransgenic Res 1889ndash97 2009

c Insertion of the ACC deaminase gene


The gene is obtained from Pseudomonas chlororaphis

(a common nonpathogenic soil bacterium)

It converts ACC to a different compound

Reduce the amount of ACC available for ethylene production

90-97 reduced ethylene production

Klee et al1991

Plants transformed with ACC

deaminase

No differences in softness

Major difference in degradation

of fruit that occurs following

ripening

Klee et al Ripening Physiology of Fruit from Transgenic Tomato (Lycopersicon

esculentum) Plants with Reduced Ethylene Synthesis Plant Physiol Vol 102 1993



d Insertion of the SAM hydrolase gene

The gene is obtained from E coli T3 bacteriophage

SAM is converted to homoserine

The amount of its precursor metabolite is reduced

Matto 2002

Good et al 1994


Regulation of Cell wall degradation

aPolygalacturonase (PG)

degrades pectin

Antisense RNA techniques

The transgenic fruit with decreased levels of PG activity

1)Do not get overly soft when ripe

2)Show less damage due to fungal infection and

3)Have elevated levels of soluble solids

Bird et al 1988

Chimaeric polygalacturonase (PG)

gene

Produce a truncated PG transcript

constitutively

Expression of the endogenous PG

gene was inhibited


CJS Smith et al Expression of a truncated tomato polygalacturonase gene inhibits

expression of the endogenous gene in transgenic plants Mol Gen Genet224477-481

1999

bPectin methylesterase (PME)

Involved in metabolism of pectin

Break large polymers into shorter molecules

Antisense RNA approach

Transgenic fruit resulted in reduced pectin depolymerization

However there was no effect on firmness during ripening

Tieman et al1992

Hall et al1993


cβ-galactosidaseNormally upregulated during the early stages of ripening

Serves to remove pectic galactan side chains

Antisense regulation

dPhospholipase DHydrolyze phospholipids

An antisense phospholipase D (PLD) cDNA Construct

resulted in a 30-40 reduction of PLD activity in ripe fruits

Transgenic fruits were firmer possessed better red colour and flavour

Pinhero et al 2003

eDeoxyhypusine synthaseAntisense gen copy of Senescence-induced deoxyhypusine synthase and

senescence-induced elf-5a

Pleiotropic effects on growth and development of tomato

Transgenics ripened normally but exhibited delayed postharvest softening

Wang et al 2005


Control of Ethylene Perception and signaling

Modifying ethylene receptors

The gene ETR1 encode an ethylene binding protein

Modified ETR1 lack the ability to respond to ethylene

Down-regulate specific tomato ethylene receptor isoforms using antisense

suppression have been reported for SlETR1 NR and SlETR4

Reporter genes related to ethylene responses and fruit ripening LeCTR1 and

SlEILs genes were also successfully silenced

Fu et al 2005

Zhu et al 2006


Lucille Alexander Journal of Experimental Botany Vol 53 No 377JC Stearns BR Glick Biotechnology Advances 21 (2003) 193ndash210

Anthocyanins Double the Shelf Life of

Tomatoes by Delaying Over ripening

Binding of specific trans-acting factors to the cognate cis-elements

Governs the spatial and temporal expression of a number of inducible genes

Tomato

E8 (Deikman et al 1998)

2A11 (Vanand Houck 1993)

Apple

ACO (Atkinson et al1998)

Melon

cucumisin (Yamagata et al 2002)

WSP (Wu et al 2003)

Strawberry

GalUR (Agius et al2005)

Grape

VvAlb1 (Li and Gray 2005)

Banana

MaExp1 (Trivedi and Nath 2004)

Fruit specific and ripening related

promoterscis-elements

Research in Environment and Life Sciences 2008

Advantages of Delayed fruit ripening

Assurance of top quality

Allowing the fruits to exude full quality

Consumers will get value for their money

Widening of market opportunities

Reduction in postharvest losses

httpwwwisaaaorgkc

References

James J Giovannoni Genetic Regulation of Fruit Development and Ripening The

Plant Cell Vol 16 S170ndashS180 2004

Antonio J Matas et al Biology and genetic engineering of fruit maturation for enhanced

quality and shelf-life Current Opinion in Biotechnology 20197ndash203 2009

V Prasanna et al Fruit Ripening PhenomenandashAn Overview Critical Reviews in Food

Science and Nutrition 471ndash19 2007

M Bouzayen et al Mechanism of Fruit Ripening Open Archive TOULOUSE Archive

Ouverte Eprints ID 4525

Aide Wang et al Null Mutation of the MdACS3 Gene Coding for a Ripening-Specific

1-Aminocyclopropane-1-Carboxylate Synthase Leads to Long Shelf Life in Apple Fruit

Plant Physiology Vol 151 pp 391ndash399 2009

Rodolfo Loacutepez-Goacutemez et al Ripening in papaya fruit is altered by ACC oxidase

Cosuppression Transgenic Res 1889ndash97 2009

Aarti Gupta et al Delayed ripening and improved fruit processing quality in tomato by

RNAi-mediated silencing of three homologs of 1-aminopropane-1-carboxylate

synthase gene Journal of Plant Physiology 170987ndash 995 2013

Liu C et al Cloning of 1-aminocyclopropane-1-carboxylate (ACC) synthetase cDNA

and the inhibition of fruit ripening by its antisense RNA in transgenic tomato plants Chin

J Biotechnol 199814(2)75-84

Gray J et al Molecular biology of fruit ripening and its manipulation

with antisense genes Plant Mol Biol 1992 May19(1)69-87

Oeller PW et al Reversible inhibition of tomato fruit senescence by antisense RNA

Science 1991 Oct 18254(5030)437-9

Harpster MH Constitutive overexpression of a ripening-related pepper endo-14-beta-

glucanase in transgenic tomato fruit does not increase xyloglucan depolymerization or

fruit softening Plant Mol Biol 2002 Oct50(3)357-69

Brummell DA et al Cell wall metabolism in fruit softening and quality and its

manipulation in transgenic plants Plant Mol Biol 2001 Sep47(1-2)311-40

Websiteshttpagbiosafetyunleduflashantisenseswf

httpwwwisaaaorgkc

httpwwwukessayscom essaysbiologyquality-and-shelf-life-of-fruits-and-

vegetablesphp

httpshodhgangainflibnetacinbitstream1060340711616_referencespdf

BooksBiology and biotechnology of the plant hormone ethylene

Edited by- A Khanellis

Transgenic plants and crops

Edited by- M Dekkerlne

References

THANK YOU

Highly coordinated

Genetically programmed

Irreversible phenomenon

Physiological biochemical changes

Development of a soft and edible ripe fruit

What is Fruit Ripening














during ripening

VPrasanna et al



Rate limiting step


The expression of


and ethylene

perception genes



Kevany et al 2007




parenchymatus cells



35 pectin

25 cellulose

20 hemicellulose














Simple genetics







and ripening



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU














during ripening

VPrasanna et al



Rate limiting step


The expression of


and ethylene

perception genes



Kevany et al 2007




parenchymatus cells



35 pectin

25 cellulose

20 hemicellulose














Simple genetics







and ripening



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU


Rate limiting step


The expression of


and ethylene

perception genes



Kevany et al 2007




parenchymatus cells



35 pectin

25 cellulose

20 hemicellulose














Simple genetics







and ripening



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU

The expression of


and ethylene

perception genes



Kevany et al 2007




parenchymatus cells



35 pectin

25 cellulose

20 hemicellulose














Simple genetics







and ripening



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU




parenchymatus cells



35 pectin

25 cellulose

20 hemicellulose














Simple genetics







and ripening



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU








Simple genetics







and ripening



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU



Labor intensive

costly








-mushy and watery


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU


vegetables







Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU

Transgenic approach

Delayed fruit

ripening











Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU







Oeller et al 1991

Yao et al1999

Nath et al 2006







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU







Aide Wang 2009



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU



homologs









Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU






Hamilton et al 1990

Ye et al 1996

Xiong et al 2003






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU






terminator










Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU








Klee et al1991


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU


deaminase




ripening









Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU






Matto 2002

Good et al 1994




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU




degrades pectin






Bird et al 1988


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU


gene


constitutively


gene was inhibited




1999







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU







Tieman et al1992

Hall et al1993









Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU








Pinhero et al 2003





Wang et al 2005










Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU









Fu et al 2005

Zhu et al 2006







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU







Tomato



Apple


Melon


WSP (Wu et al 2003)

Strawberry


Grape


Banana











httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU







httpwwwisaaaorgkc

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU

References























Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU


Science 1991 Oct 18254(5030)437-9







httpwwwisaaaorgkc


vegetablesphp






References

THANK YOU

THANK YOU

transgenics for delayed fruit ripening

Science

ethylene productionwhat

ethylene perception

tomato fruit development

yao et

nath et

ripeningthe plant cell

low expression of ripening

synthase geneoeller