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coffee plant and seed but also in other plants (Douglas, 1996;Grace et al., 1998; Materska and Perucka, 2005; Pennycookeet al., 2005). The use of nitrogen-rich fertilizers (Malta et al.,2003) and situations of boron deficiency (Camacho-Cristobal,et al., 2002) have also shown to increase the content of totalCGA in coffee seeds and other plants.Small amounts of phenolic compounds differentfrom CGA and related compounds have been identifiedin green coffee beans. The presence of about 1 % ofphenolic glycosides has been suggested (Clifford, 1985).Anthocyanidins such as cyanidins, pelargonidins and onepeonidin were identified in arabica coffee seeds as a residueof red skin fruits. (Mazza and Miniati, 1993). Lignanssuch as secoisolariciresinol, lariciresinol, matairesinoland pinoresinol, in order of abundance, are also presentin coffee (Milder et al., 2005). Some free volatile phenolshave been identified (Moreira et al., 2000). Tannins havenot been identified in wet processed coffee seeds (Cliffordand Ramirez-Martinez, 1991a). However, it is possible thatdry processed beans contain a small amount of tannins as aresidue of the coffee pulp.Biosynthesis of chlorogenic acids and relevance for theplant physiologyCGA are products of the phenylpropanoid pathway, onebranch of the phenolic metabolism in higher plants that isinduced in response to environmental stress conditions suchas infection by microbial pathogens, mechanical wounding,and excessive UV or high visible light levels (Herrmann,1995; Haard and Chism, 1996). Plant phenolic acidsare synthesized from phenylalanine and tyrosine via theshikimic acid pathway, which converts simple carbohydrateprecursors, derived from glycolysis and the pentosephosphate shunt (phospho-enol-pyruvate and D-erythrose-4-phosphate), into aromatic amino acids (figure 2). Theparent trans-cinnamic acid is formed from L-phenylalanineby the action of phenylalanine ammonia-lyase, a keyenzyme in the biosynthesis of phenolic compounds that isactivated in response to different stress conditions. Quinicacid is synthesized from 3-dehydroquinate, an intermediatemetabolite of the shikimic acid pathway. Hydroxy-cinnamicacids (p-coumaric, caffeic, ferulic and sinapic) may besynthesized from cinnamic acid by hydroxylation oralternatively from tyrosine by the action of tyrosine aminolyase(figure 2).The final steps of biosynthesis of the 5-monoacyl CGA(5-CQA, 5-FQA and 5-p-CoQA) have been studied in severalplants as well as coffee and appear to include binding oftrans-cinnamic acid to coenzyme A (CoA) by a CoA lyase,followed by transfer to quinic acid by a cinnamoyl transferase(Gross, 1981). The origin of the CGA with acyl groups inpositions 3- and 4- is unclear, although the possibility of acylmigration has been considered (Gross, 1981). Once formed,phenolic acids and CGA can be substrate of enzymes suchas polyphenol oxidase (Mazzafera and Robinson, 2000) and
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