Pectin is an important polysaccharide with applications in foods, Pharmaceuticals, and a number of other industries. Its importance in the food sector lies in its ability to form gel in the presence of Ca2+ ions or a solute at low pH. Although the exact mechanism of gel formation is not clear, significant progress has been made in this direction. Depending on the pectin, coordinate bonding with Ca2+ ions or hydrogen bonding and hydrophobic interactions are involved in gel formation. In low‐methoxyl pectin, gelation results from ionic linkage via calcium bridges between two carboxyl groups belonging to two different chains in close contact with each other. In high‐methoxyl pectin, the cross‐linking of pectin molecules involves a combination of hydrogen bonds and hydrophobic interactions between the molecules. A number of factors—pH, presence of other solutes, molecular size, degree of methoxylation, number and arrangement of side chains, and charge density on the molecule— influence the gelation o...

Chemistry and uses of pectin--a review.

Excessive softening is the main factor limiting fruit shelf life and storage. Transgenic plants modified in the expression of cell wall modifying proteins have been used to investigate the role of particular activities in fruit softening during ripening, and in the manufacture of processed fruit products. Transgenic experiments show that polygalacturonase (PG) activity is largely responsible for pectin depolymerization and solubilization, but that PG-mediated pectin depolymerization requires pectin to be de-methyl-esterified by pectin methylesterase (PME), and that the PG β-subunit protein plays a role in limiting pectin solubilization. Suppression of PG activity only slightly reduces fruit softening (but extends fruit shelf life), suppression of PME activity does not affect firmness during normal ripening, and suppression of β-subunit protein accumulation increases softening. All these pectin-modifying proteins affect the integrity of the middle lamella, which controls cell-to-cell adhesion and thus influences fruit texture. Diminished accumulation of either PG or PME activity considerably increases the viscosity of tomato juice or paste, which is correlated with reduced polyuronide depolymerization during processing. In contrast, suppression of β-galactosidase activity early in ripening significantly reduces fruit softening, suggesting that the removal of pectic galactan side-chains is an important factor in the cell wall changes leading to ripening-related firmness loss. Suppression or overexpression of endo-(1\to4)β-d-glucanase activity has no detectable effect on fruit softening or the depolymerization of matrix glycans, and neither the substrate nor the function for this enzyme has been determined. The role of xyloglucan endotransglycosylase activity in softening is also obscure, and the activity responsible for xyloglucan depolymerization during ripening, a major contributor to softening, has not yet been identified. However, ripening-related expansin protein abundance is directly correlated with fruit softening and has additional indirect effects on pectin depolymerization, showing that this protein is intimately involved in the softening process. Transgenic work has shown that the cell wall changes leading to fruit softening and textural changes are complex, and involve the coordinated and interdependent activities of a range of cell wall-modifying proteins. It is suggested that the cell wall changes caused early in ripening by the activities of some enzymes, notably β-galactosidase and ripening-related expansin, may restrict or control the activities of other ripening-related enzymes necessary for the fruit softening process.

Cell wall metabolism in fruit softening and quality and its manipulation in transgenic plants

https://ucanr.edu/datastoreFiles/608-360.pdf

Biochemical bases of appearance and texture changes in fresh-cut fruit and vegetables

Fruit softening during ripening involves a coordinated series of modifications to the polysaccharide components of the primary cell wall and middle lamella, resulting in a weakening of the structure. Degradation of polysaccharides and alterations in the bonding between polymers cause an increase in cell separation and a softening and swelling of the wall, which, combined with alterations in turgor, bring about fruit softening and textural changes. A wide range in the extent of cell wall pectic modifications has been observed between species, whereas the depolymerisation of xyloglucan is relatively limited and more consistent. The earliest events to be initiated are usually a loss of pectic galactan side chains and the depolymerisation of matrix glycans, which may begin before ripening, followed by a loss of pectic arabinan side chains and pectin solubilisation. The depolymerisation of pectins may begin during early to mid-ripening, but is usually most pronounced late in ripening. However, some of these events may be absent or occur at very low levels in some species. Cell wall swelling may be related to a loosening of the xyloglucan-cellulose network and to pectin solubilisation, and these processes combined with the loss of pectic side chains increase wall porosity. An increase in wall porosity later in ripening may allow increased access of degradative enzymes to their substrates.

Cell wall disassembly in ripening fruit

Fruits constitute a commercially important and nutritionally indispensable food commodity. Being a part of a balanced diet, fruits play a vital role in human nutrition by supplying the necessary growth regulating factors essential for maintaining normal health. Fruits are widely distributed in nature. One of the limiting factors that influence their economic value is the relatively short ripening period and reduced post-harvest life. Fruit ripening is a highly coordinated, genetically programmed, and an irreversible phenomenon involving a series of physiological, biochemical, and organoleptic changes, that finally leads to the development of a soft edible ripe fruit with desirable quality attributes. Excessive textural softening during ripening leads to adverse effects/spoilage upon storage. Carbohydrates play a major role in the ripening process, by way of depolymerization leading to decreased molecular size with concomitant increase in the levels of ripening inducing specific enzymes, whose target differ from fruit to fruit. The major classes of cell wall polysaccharides that undergo modifications during ripening are starch, pectins, cellulose, and hemicelluloses. Pectins are the common and major components of primary cell wall and middle lamella, contributing to the texture and quality of fruits. Their degradation during ripening seems to be responsible for tissue softening of a number of fruits. Structurally pectins are a diverse group of heteropolysaccharides containing partially methylated D-galacturonic acid residues with side chain appendages of several neutral polysaccharides. The degree of polymerization/esterification and the proportion of neutral sugar residues/side chains are the principal factors contributing to their (micro-) heterogeneity. Pectin degrading enzymes such as polygalacturonase, pectin methyl esterase, lyase, and rhamnogalacturonase are the most implicated in fruit-tissue softening. Recent advances in molecular biology have provided a better understanding of the biochemistry of fruit ripening as well as providing a hand for genetic manipulation of the entire ripening process. It is desirable that significant breakthroughs in such related areas will come forth in the near future, leading to considerable societal benefits.

Fruit Ripening Phenomena–An Overview

Polyuronides and hemicelluloses derived from ethanol powders or cell walls of strawberry (Fragaria × ananassa, Duch. Dover) receptacle tissue were examined to determine if these wall polymers might be involved in the softening of this fruit. Throughout maturation and ripening, total polyuronides increased on a per fruit basis although as a percentage of ethanol powder they remained constant. Gel-filtration chromatography confirmed that polyuronide solubility was not correlated with extensive enzymic hydrolysis, an observation consistent with the fact that D-galacturonanase (polygalacturonase) activity was not detected in strawberry fruit. The sugar composition of alkali-soluble wall polymers showed little change throughout development. However, changes occurred in the molecular weights of these polymers during ripening.

D. J. Huber

Papers

Strawberry Fruit Softening: The Potential Roles of Polyuronides and Hemicelluloses