Biology and genetic engineering of fruit maturation for enhanced quality and shelf-life
Section snippets
Ethylene and fruit ripening
Ripening is regulated by both internal and external stimuli, including temperature, light, plant nutrient status, water availability, and hormones. Many fruits, in particular the so-called climacteric fruits, require ethylene for ripening, resulting in the targeting of this hormone as a means of ripening control. Although nonclimacteric fruits typically produce little ethylene during ripening, many have still been shown to be affected by exogenous ethylene during ripening, making ethylene
Ethylene response genes: potential targets for ethylene response and ripening control
Ethylene signal transduction genes have been well characterized in Arabidopsis (for recent reviews see [4, 5, 6]). More recent efforts in understanding the ethylene response during fruit ripening have focused on the characterization of tomato homologs. All the components of the Arabidopsis ethylene signal transduction analyzed thus far are conserved in tomato (reviewed in [2]). However, the family size and expression profiles of some of these genes differ between Arabidopsis and tomato. For
Transcriptional control of fruit ripening and conserved ripening regulators
Although considerable effort has focused on ethylene synthesis and response, only recently have inroads been made into understanding ripening control before ethylene: a regulatory system that is possibly conserved between climacteric and nonclimacteric species. Recently, the mutated genes underlying the ripening mutants RIPENING INHIBITOR (RIN) and COLOURLESS NON RIPENING (CNR) were cloned, providing key insights into the regulation of the ripening process upstream of ethylene. The rin and Cnr
Genetic manipulation of ripening regulatory genes
To date, attempts to manipulate the expression of target genes during fruit ripening have produced varying results. As mentioned, transcriptional control has been utilized with respect to breeding for heterozygosity at the rin mutation (Rin/rin) in commercial lines to delay ripening and extend shelf-life; however, this modification leads to the inhibition of flavor and nutritional compound accumulation along with undesirable textural traits in some backgrounds due to incomplete ripening. In
Cell walls and cuticles as key factors for fruit shelf-life
The cell wall has a profoundly important influence on fruit texture and cell wall components and the underlying genes have been frequent targets for genetic engineering, mostly in tomato, with the goal of extending shelf-life [19, 20]. Although knowledge of the mechanism of fruit softening has grown in recent years, it has proven especially difficult to establish the relationship between specific aspects of cell wall metabolism or architecture and their relationship to changes in tissue
When a fruit is more than a fruit?
Fresh fruits and their processed derivates represent important sources of carotenoids, flavonoids, and anthocyanins and the possibility of improving content using bioengineering represent an opportunity to improve access to healthy foods. The inherent resiliency of plant metabolism to maintain homeostasis has impaired efforts to facilitate changes in some of these pathways, as has been especially well documented for carotenoids [29]. An alternate approach is to focus on regulatory rather than
New tools, new strategies, and enhancement of traditional approaches
Prior efforts toward fruit quality improvement have focused on single steps in biochemical pathways, but complex feedback regulation networks have complicated such strategies, as noted above. Targeting regulatory genes may provide a route to avoid these regulatory impediments though the difficult work of assessing the impact of altering specific pathway steps is required to understand the underlying regulatory networks [32]. The use of a transcription factor that regulates a complete
Prospects for the future
It is important to note that recent studies have reported that genetically modified organisms are not inherently dangerous and can be considered safe to consumers when compared with conventional foods developed for target traits through naturally occurring genetic variation [49]. Of course, given that the safety of crops is a primary concern for both producers and consumers, it is necessary to define the risk and unintended effects in genetically modified plants [50], including the effects of a
References and recommended reading
Papers of particular interest, published within the period of review, have been highlighted as:
• of special interest
•• of outstanding interest
Acknowledgements
We acknowledge the support from USDA-NRI grants 2007-02773 and 2006-35304-17323, and the NSF Plant Genome grants 05-01778 and 06-06595. Support was also provided by a grant to JKCR from the US–Israel Binational Science Foundation, and to NEG from the New Zealand Foundation for Research, Science and Technology FRST-BTPR0701.
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These authors contributed equally to this manuscript and should be considered co-first authors.