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              Biopolymers                                                                                 227

                                                                  In many plant cell walls, the major hemicellulose is ei-
                                                                ther arabinoxylan—main chains of xylose with branches
                                                                of single arabinose residues on some of the xylose rings—
                                                                or glucuronoarabinoxylans with additional branches of
                                                                single unsubstituted or modified glucuronic acid residues
                                                                on some xylose rings.
                                                                  Xyloglucans may act as food reserves in seeds and have
                                                                a structural role in mature plants. They probably have ex-
                                                                tended ribbon-like structures. Xyloglucans have backbone
                                                                chains of β-(1 → 4)-linked glucose residues with xylose
                                                                units linked α-(1 → 6) as branches on some of the glucose
                                                                rings. Some branches can also contain galactose and fu-
                                                                cose residues but there seems to be no regular repeat in the
                                                                structure. This is a feature of many plant polysaccharides,
                                                                and the apparent lack of regularity may be necessary for
                                                                interaction with other proteins and polysaccharides in cell
                                                                walls which can, as a plant grows, elongate to many times
                                                                their original size. In addition, arabinoxyloglucans exist
                                                                in some plants, with arabinose units linked α-(1 → 2) to
                                                                xylose.
                                                                  Glucomannans and the related galactoglucomannans
                                                                are important in softwood, and do not have a regular struc-
                                                                ture but have linear main chains containing, in an appar-
                                                                ently random order, both mannose and glucose joined
              FIGURE 12 Representation of calcium pectate gels with
                                                                by β-(1→4)-links. The galactoglucomannans have, in
              “egg box” model for chain associations of nonesterified α-D-
                                                                addition, single-residue branches of galactose joined by
              galacturonan regions through calcium ions, interrupted by ester-
              ified α-D-galacturonan regions, or terminated by insertion of L-  α-(1 → 6)-bonds to some mannose units. Related polysac-
              rhamnose residues in the chain. [Adapted with permission from  charides are the galactomannans which have food reserve
              Rees, D. A., Morris, E. R., Thom, D., and Madden, J. K. (1982). In  and structural functions in plants. These have backbone
              “The Polysaccharides”, (G. O. Aspinall, ed.), Vol. 1, p. 265. Copy-  chains of β-(1 → 4)-linked mannose residues, with single-
              right 1982 by Academic Press, Inc., New York. From Aspinall,
              G. O. “Polysaccharides” in the Encyclopedia of Physical Science  residue branches of galactose joined by α-(1 → 6)-bonds
              and Technology, Vol. 11, p. 184. Copyright 1987 by Academic  to some mannose units. Two of these galactomannans are
              Press, Inc., New York.]                           commercially important. They are guar gum and locust
                                                                bean (also known as carob) gum, isolated from the en-
                                                                dosperm of seeds of the leguminous plants Cyamopsis
              structure where chains associate to form junction zones  tetragonolobus and Ceratonia siliqua, respectively. The
              (see  Figs.  10c  and  12).  Because  high  sucrose  concen-  guar gum has a higher content of galactose than the locust
              trations are not needed for gel formation, low-methoxyl  bean gum and is more water soluble. These galactoman-
              pectins can be used for low-sugar jams and jellies. En-  nans give high-viscosity solutions in water and so are used
              zymic degradation of pectins in cell walls is necessary for  as thickening agents in the food industry and in the phar-
              the softening of fruit during the ripening process. Apple  maceutical industry. In particular, they give good gels with
              juices must be clarified by enzymic degradation of pectins  carrageenans, agarose, or xanthan (see next sections) and
              in freshly squeezed juice; in contrast the pectin-modifying  can be used in ice creams and cheese spreads. The galac-
              enzymes of orange juice must be deactivated by storing the  tose side groups are believed to interrupt junction zones
              juice frozen, in order to maintain the cloudy appearance  in the gels. Derivatives of locust bean gum are of use in
              of the juice.                                     the textile industry as sizes, while guar gum can act as
                The polysaccharides generally considered as hemicel-  a dye thickener. Guar gum also increases the strength of
              luloses have backbones composed of xylose residues, as  cellulosic fiber in paper-making.
              in the xylans, or glucose units, as in the xyloglucans, or  Matrix polysaccharides, mainly the pectins and hemi-
              mannose sometimes in combination with glucose, as in  celluloses,ofplant material,e.g., cereals, fruit,and vegeta-
              the mannans and glucomannans. Few of these are isolated  bles, in the human diet are termed dietary fiber, although
              for commercial exploitation.                      they may be more gel-like than fibrous. These polymers
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