Changes in pericarp morphology, physiology and cell wall composition account for flesh firmness during the ripening of blackberry (Rubus spp.) fruit

Abstract

Fruit flesh firmness plays a critical role in controlling blackberry (Rubus spp.) postharvest shelf life. This work aimed to identify the underlying characteristics in pericarp morphology, cell wall hydrolase activity and cell wall composition that account for flesh firmness in soft-fruited ‘Boysen’ and firm-fruited ‘Arapaho’ cultivars that have a similar ripening time of 39 days after flowering (DAF). The fruit firmness of both fruits decreased noticeably at the onset of color change, and this decrease hastened from 33 DAF to ripening despite considerable differences in their temporal changes. The evaluation of pericarp cellular morphology revealed that the disassembly of the cell wall in both fruits was likely initiated at 33 DAF, followed by extreme degradation at 39 DAF. Cell wall hydrolase activity assays indicated that increases in polygalacturonase (PG) and cellulase activity also dramatically occurred in the late softening stages of both fruits. Notably, appreciably higher levels of cellulase, and significant increases in pectin methylesterase (PME), α-L-arabinofuranosidase (α-L-Af) and xyloglucan endotransglycosylase (XET) were only detected in the late ripening stages of ‘Boysen’. In terms of cell wall components, the levels of cell wall material (CWM), cellulose and hemicellulose declined similarly during ripening in both fruits. Comparatively, lower levels of CWM, chelator soluble pectin (CSP), sodium carbonate soluble pectin (SSP), and hemicellulose as well as higher levels of water soluble pectin (WSP) were found in ‘Boysen’ than those in ‘Arapaho’ during maturation and ripening. Overall, the loss of fruit firmness during ripening in blackberry fruit is correlated closely with biochemical changes in cell wall fractions that involve hydrolytic processes, resulting in the breakdown of cell-wall polymers. The low firmness of ‘Boysen’, as determined by cell wall degradation, could be of particular relevance to the function of cell wall degrading enzymes and to the more drastic degradation of cell wall components than that in the firm cv. ‘Arapaho’.