amylopectin

Summary

Summary: A highly branched glucan in starch.

Top Publications

  1. Denyer K, Waite D, Edwards A, Martin C, Smith A. Interaction with amylopectin influences the ability of granule-bound starch synthase I to elongate malto-oligosaccharides. Biochem J. 1999;342 Pt 3:647-53 pubmed
    ..One of these, granule-bound starch synthase I (GBSSI), is responsible for the synthesis of amylose inside the amylopectin matrix of the starch granule in vivo...
  2. Fujita N, Yoshida M, Asakura N, Ohdan T, Miyao A, Hirochika H, et al. Function and characterization of starch synthase I using mutants in rice. Plant Physiol. 2006;140:1070-84 pubmed
    ..The mutant endosperm amylopectin showed a decrease in chains with degree of polymerization (DP) 8 to 12 and an increase in chains with DP 6 to 7 ..
  3. Engelsen S, Madsen A, Blennow A, Motawia M, Møller B, Larsen S. The phosphorylation site in double helical amylopectin as investigated by a combined approach using chemical synthesis, crystallography and molecular modeling. FEBS Lett. 2003;541:137-44 pubmed
    ..and through modeling extrapolate the results to the double helical structure of the crystalline part of amylopectin. The geometries of the existing crystal structures of 6-O-phosphate groups were found to belong to two main ..
  4. Lloyd J, Landschütze V, Kossmann J. Simultaneous antisense inhibition of two starch-synthase isoforms in potato tubers leads to accumulation of grossly modified amylopectin. Biochem J. 1999;338 ( Pt 2):515-21 pubmed
    ..In order to understand why the starch granules were distorted, amylopectin was isolated and the constituent chain lengths analysed...
  5. Syahariza Z, Sar S, Hasjim J, Tizzotti M, Gilbert R. The importance of amylose and amylopectin fine structures for starch digestibility in cooked rice grains. Food Chem. 2013;136:742-9 pubmed publisher
    ..characteristics, including fine structures of the distributions of branch (chain) lengths in both amylose and amylopectin. The in vitro digestion rate tends to increase with longer amylose branches and smaller ratios of long ..
  6. Zhang X, Szydlowski N, Delvalle D, D Hulst C, James M, Myers A. Overlapping functions of the starch synthases SSII and SSIII in amylopectin biosynthesis in Arabidopsis. BMC Plant Biol. 2008;8:96 pubmed publisher
    The biochemical mechanisms that determine the molecular architecture of amylopectin are central in plant biology because they allow long-term storage of reduced carbon...
  7. Hansen P, Larsen F, Motawia S, Blennow A, Spraul M, Dvortsak P, et al. Structure and hydration of the amylopectin trisaccharide building blocks--Synthesis, NMR, and molecular dynamics. Biopolymers. 2008;89:1179-93 pubmed publisher
    To gain insight into the molecular details and hydration of amylopectin, the five constituting trisaccharides have been chemically synthesized as their methyl alpha-glycosides...
  8. Hannah L, James M. The complexities of starch biosynthesis in cereal endosperms. Curr Opin Biotechnol. 2008;19:160-5 pubmed publisher
    ..of starch is the origin of nonrandom or clustered alpha-1,6 branch-points within the major component of starch, amylopectin. Developing evidence that several of the starch biosynthetic enzymes involved in amylopectin synthesis occur in ..
  9. Delatte T, Trevisan M, Parker M, Zeeman S. Arabidopsis mutants Atisa1 and Atisa2 have identical phenotypes and lack the same multimeric isoamylase, which influences the branch point distribution of amylopectin during starch synthesis. Plant J. 2005;41:815-30 pubmed
    ..The amylopectin of the remaining starch and the phytoglycogen in the mutants are structurally related to each other and differ ..

More Information

Publications62

  1. Stahl Y, Coates S, Bryce J, Morris P. Antisense downregulation of the barley limit dextrinase inhibitor modulates starch granule size distribution, starch composition and amylopectin structure. Plant J. 2004;39:599-611 pubmed
    ..starch showed much reduced numbers of the small B-type starch granules, as well as reduced amylose relative to amylopectin levels and reduced total starch...
  2. Lee S, Nilsson P, Nilsson G, Wahlund K. Development of asymmetrical flow field-flow fractionation-multi angle laser light scattering analysis for molecular mass characterization of cationic potato amylopectin. J Chromatogr A. 2003;1011:111-23 pubmed
    ..AsFlFFF)-multi angle laser light scattering (MALLS), and to develop a method for analysis of cationic potato amylopectin (CPAP) having ultrahigh molecular mass (UHMr)...
  3. James M, Denyer K, Myers A. Starch synthesis in the cereal endosperm. Curr Opin Plant Biol. 2003;6:215-22 pubmed
    ..For the first time, tools for global analyses of starch biosynthesis are available for cereal crops, and are heralded by the draft sequence of the rice genome. ..
  4. Zhu P, Tsang R, Tsai C. Nonencapsulated Neisseria meningitidis strain produces amylopectin from sucrose: altering the concept for differentiation between N. meningitidis and N. polysaccharea. J Clin Microbiol. 2003;41:273-8 pubmed
    ..Neisseria polysaccharea is a nonpathogenic species. N. polysaccharea is able to use sucrose to produce amylopectin, a starch-like polysaccharide, which distinguishes it biochemically from the pathogenic species N. meningitidis...
  5. Fulton D, Edwards A, Pilling E, Robinson H, Fahy B, Seale R, et al. Role of granule-bound starch synthase in determination of amylopectin structure and starch granule morphology in potato. J Biol Chem. 2002;277:10834-41 pubmed
    Reductions in activity of SSIII, the major isoform of starch synthase responsible for amylopectin synthesis in the potato tuber, result in fissuring of the starch granules...
  6. Jiang H, Dian W, Wu P. Effect of high temperature on fine structure of amylopectin in rice endosperm by reducing the activity of the starch branching enzyme. Phytochemistry. 2003;63:53-9 pubmed
    ..The activities and gene expression of key enzymes for the biosynthesis of amylose and amylopectin were examined...
  7. Maddelein M, Libessart N, Bellanger F, Delrue B, D Hulst C, Van Den Koornhuyse N, et al. Toward an understanding of the biogenesis of the starch granule. Determination of granule-bound and soluble starch synthase functions in amylopectin synthesis. J Biol Chem. 1994;269:25150-7 pubmed
    ..has precluded genetic assignment of functions to the various soluble starch synthases in the building of amylopectin. In Chlamydomonas, we have recently shown that defects in the major soluble starch synthase lead to a specific ..
  8. Fujita N, Kubo A, Suh D, Wong K, Jane J, Ozawa K, et al. Antisense inhibition of isoamylase alters the structure of amylopectin and the physicochemical properties of starch in rice endosperm. Plant Cell Physiol. 2003;44:607-18 pubmed
    This is the first report on regulation of the isoamylase1 gene to modify the structure of amylopectin and properties of starch by using antisense technology in plants...
  9. Nishi A, Nakamura Y, Tanaka N, Satoh H. Biochemical and genetic analysis of the effects of amylose-extender mutation in rice endosperm. Plant Physiol. 2001;127:459-72 pubmed
    ..that the mutation in the gene for starch-branching enzyme IIb (BEIIb) specifically altered the structure of amylopectin in the endosperm by reducing short chains with degree of polymerization of 17 or less, with the greatest ..
  10. Bluhm B, Woloshuk C. Amylopectin induces fumonisin B1 production by Fusarium verticillioides during colonization of maize kernels. Mol Plant Microbe Interact. 2005;18:1333-9 pubmed
    ..the wild-type strain produced only trace amounts of FB1, but it produced large amounts of FB1 when grown on amylopectin or dextrin, a product of amylopectin hydrolysis. We conclude that amylopectin induces FB1 production in F...
  11. Blennow A, Mette Bay Smidt A, Bauer R. Amylopectin aggregation as a function of starch phosphate content studied by size exclusion chromatography and on-line refractive index and light scattering. Int J Biol Macromol. 2001;28:409-20 pubmed
    ..Three major regions in the SEC profile were identified, consisting of large amylopectin aggregates, amylopectin particles with radius of gyration (Rg) of approx 200 nm (400 nm blocklets) and amylose...
  12. Zhang X, Colleoni C, Ratushna V, Sirghie Colleoni M, James M, Myers A. Molecular characterization demonstrates that the Zea mays gene sugary2 codes for the starch synthase isoform SSIIa. Plant Mol Biol. 2004;54:865-79 pubmed
    ..structural effect of the su2 (-) mutations was shown to be increased abundance of short glucan chains in amylopectin and a proportional decrease in intermediate length chains, similar to the effects of SSII deficiency in other ..
  13. Hanashiro I, Tagawa M, Shibahara S, Iwata K, Takeda Y. Examination of molar-based distribution of A, B and C chains of amylopectin by fluorescent labeling with 2-aminopyridine. Carbohydr Res. 2002;337:1211-5 pubmed
    A method for determination of a molar-based distribution of A, B and C chains of amylopectin was developed...
  14. Lu B, Guo Z, Liang J. Effects of the activities of key enzymes involved in starch biosynthesis on the fine structure of amylopectin in developing rice (Oryza sativa L.) endosperms. Sci China C Life Sci. 2008;51:863-71 pubmed publisher
    ..branching enzyme (SBE) and starch debranching enzymes (DBE) were studied, and changes of fine structure of amylopectin were characterized by isoamylase treatment during rice grain development, using trans anti-waxy gene rice ..
  15. Zeeman S, Kossmann J, Smith A. Starch: its metabolism, evolution, and biotechnological modification in plants. Annu Rev Plant Biol. 2010;61:209-34 pubmed publisher
    ..First, we assess progress in identifying the enzymatic machinery required for the synthesis of amylopectin, the glucose polymer responsible for the insoluble nature of starch...
  16. Hanashiro I, Itoh K, Kuratomi Y, Yamazaki M, Igarashi T, Matsugasako J, et al. Granule-bound starch synthase I is responsible for biosynthesis of extra-long unit chains of amylopectin in rice. Plant Cell Physiol. 2008;49:925-33 pubmed publisher
    ..The increase was in part due to a significant amount of extra-long unit chains (ELCs) of amylopectin (7.5-8.4% of amylopectin weight), that were absent in the non-transgenic wx cultivars...
  17. Benmoussa M, Moldenhauer K, Hamaker B. Rice amylopectin fine structure variability affects starch digestion properties. J Agric Food Chem. 2007;55:1475-9 pubmed
    ..in rapid viscoanalyzer (RVA) pasting breakdown to study the relationship between starch digestibility and amylopectin fine structure and pasting properties...
  18. Yao Y, Thompson D, Guiltinan M. Maize starch-branching enzyme isoforms and amylopectin structure. In the absence of starch-branching enzyme IIb, the further absence of starch-branching enzyme Ia leads to increased branching. Plant Physiol. 2004;136:3515-23 pubmed
    ..the function of SBEIIb is predominant to that of SBEIa, and SBEIa would have an observable effect only on amylopectin structure in the absence of SBEIIb...
  19. Venkatesan J, Pallela R, Bhatnagar I, Kim S. Chitosan-amylopectin/hydroxyapatite and chitosan-chondroitin sulphate/hydroxyapatite composite scaffolds for bone tissue engineering. Int J Biol Macromol. 2012;51:1033-42 pubmed publisher
    ..tri-component scaffolds of chitosan/natural hydroxyapatite with chondroitin sulfate (chitosan-CS/HAp) and amylopectin (chitosan-AP/HAp) have been developed for the first time via freeze-drying method and were characterized ..
  20. Shimonaga T, Fujiwara S, Kaneko M, Izumo A, Nihei S, Francisco P, et al. Variation in storage alpha-polyglucans of red algae: amylose and semi-amylopectin types in Porphyridium and glycogen type in Cyanidium. Mar Biotechnol (NY). 2007;9:192-202 pubmed
    ..the unicellular species Porphyridium purpureum R-1 (order Porphyridiales, class Bangiophyceae) produces both amylopectin-type and amylose-type alpha-polyglucans...
  21. Utsumi Y, Nakamura Y. Structural and enzymatic characterization of the isoamylase1 homo-oligomer and the isoamylase1-isoamylase2 hetero-oligomer from rice endosperm. Planta. 2006;225:75-87 pubmed
    ..that with the homo-oligomer, although no marked differences were found in chain preferences for debranching of amylopectin and phytoglycogen between these forms...
  22. Ohdan T, Francisco P, Sawada T, Hirose T, Terao T, Satoh H, et al. Expression profiling of genes involved in starch synthesis in sink and source organs of rice. J Exp Bot. 2005;56:3229-44 pubmed
  23. Lee C, Le Q, Kim Y, Shim J, Lee S, Park J, et al. Enzymatic synthesis and properties of highly branched rice starch amylose and amylopectin cluster. J Agric Food Chem. 2008;56:126-31 pubmed
    We enzymatically modified rice starch to produce highly branched amylopectin and amylose and analyzed the resulting structural changes...
  24. Ral J, Colleoni C, Wattebled F, Dauvillee D, Nempont C, Deschamps P, et al. Circadian clock regulation of starch metabolism establishes GBSSI as a major contributor to amylopectin synthesis in Chlamydomonas reinhardtii. Plant Physiol. 2006;142:305-17 pubmed
    ..show that both enzymes play a similar function in synthesizing the long glucan fraction that interconnects the amylopectin clusters...
  25. Corzana F, Motawia M, Herve du Penhoat C, van den Berg F, Blennow A, Perez S, et al. Hydration of the amylopectin branch point. Evidence of restricted conformational diversity of the alpha-(1-->6) linkage. J Am Chem Soc. 2004;126:13144-55 pubmed
    The hydration behavior of a model compound for the amylopectin branch point, methyl 6'-alpha-maltosyl-alpha-maltotrioside, was investigated by combining molecular dynamics simulations in explicit water, 500 MHz NMR spectroscopy, ..
  26. Zhong F, Yokoyama W, Wang Q, Shoemaker C. Rice starch, amylopectin, and amylose: molecular weight and solubility in dimethyl sulfoxide-based solvents. J Agric Food Chem. 2006;54:2320-6 pubmed
    ..was used as solvent for multi-angle laser-light scattering (MALLS) batch mode analysis of rice starch, and amylopectin and amylose weight-average molecular weight (Mw)...
  27. Satoh H, Nishi A, Yamashita K, Takemoto Y, Tanaka Y, Hosaka Y, et al. Starch-branching enzyme I-deficient mutation specifically affects the structure and properties of starch in rice endosperm. Plant Physiol. 2003;133:1111-21 pubmed
    ..However, the mutation apparently altered the fine structure of amylopectin. The mutant amylopectin was characterized by significant decrease in both long chains with degree of ..
  28. Marchal L, Ulijn R, De Gooijer C, Franke G, Tramper J. Monte Carlo simulation of the alpha-amylolysis of amylopectin potato starch. 2. alpha-amylolysis of amylopectin. Bioprocess Biosyst Eng. 2003;26:123-32 pubmed
    ..Potato amylopectin, the substrate of the hydrolysis reaction, was modeled in a computer matrix...
  29. Zhou Y, Yang B, Ren X, Liu Z, Deng Z, Chen L, et al. Hyperbranched cationic amylopectin derivatives for gene delivery. Biomaterials. 2012;33:4731-40 pubmed publisher
    A series of hyperbranched cationic amylopectin derivatives conjugated with 1,2-ethylenediamine, diethylenetriamine and 3-(dimethylamino)-1-propylamine residues, named as EDA-Amp, DETA-Amp and DMAPA-Amp, were synthesized by the N,N'-..
  30. Cheng J, Zhu J, Wen N, Xiong F. Stability and pharmacokinetic studies of O-palmitoyl amylopectin anchored dipyridamole liposomes. Int J Pharm. 2006;313:136-43 pubmed
    ..b>Amylopectin was palmitoylated and anchored on the surface of plain DIP liposomes...
  31. Szydlowski N, Ragel P, Hennen Bierwagen T, Planchot V, Myers A, Mérida A, et al. Integrated functions among multiple starch synthases determine both amylopectin chain length and branch linkage location in Arabidopsis leaf starch. J Exp Bot. 2011;62:4547-59 pubmed publisher
    ..In both double mutants the residual starch was structurally modified including higher ratios of amylose:amylopectin, altered glucan chain length distribution within amylopectin, abnormal granule morphology, and altered ..
  32. Jun H, Daiwen C, Bing Y. Metabolic and transcriptomic responses of weaned pigs induced by different dietary amylose and amylopectin ratio. PLoS ONE. 2010;5:e15110 pubmed publisher
    ..However, the nutritional value of starch largely depends on its amylose and amylopectin ratio...
  33. Guerardel Y, Leleu D, Coppin A, Lienard L, Slomianny C, Strecker G, et al. Amylopectin biogenesis and characterization in the protozoan parasite Toxoplasma gondii, the intracellular development of which is restricted in the HepG2 cell line. Microbes Infect. 2005;7:41-8 pubmed
    ..their life cycle, some apicomplexan parasites accumulate a crystalline storage polysaccharide analogous to amylopectin within the cytoplasm. In T...
  34. Morell M, Kosar Hashemi B, Cmiel M, Samuel M, Chandler P, Rahman S, et al. Barley sex6 mutants lack starch synthase IIa activity and contain a starch with novel properties. Plant J. 2003;34:173-85 pubmed
    ..The loss of SSIIa activity in barley leads to novel and informative phenotypes. First, a decrease in amylopectin synthesis to less than 20% of the wild-type levels indicates that SSIIa accounts for the majority of the ..
  35. Pandey M, Rani N, Madhav M, Sundaram R, Varaprasad G, Sivaranjani A, et al. Different isoforms of starch-synthesizing enzymes controlling amylose and amylopectin content in rice (Oryza sativa L.). Biotechnol Adv. 2012;30:1697-706 pubmed publisher
    Starch, composed of amylose and amylopectin, greatly influences rice cooking and textural quality, which in turn is controlled by various isoforms of several enzymes...
  36. Delvalle D, Dumez S, Wattebled F, Roldán I, Planchot V, Berbezy P, et al. Soluble starch synthase I: a major determinant for the synthesis of amylopectin in Arabidopsis thaliana leaves. Plant J. 2005;43:398-412 pubmed
    ..These activities are involved in amylopectin synthesis and are extremely well conserved throughout the plant kingdom...
  37. Nakamura Y, Francisco P, Hosaka Y, Sato A, Sawada T, Kubo A, et al. Essential amino acids of starch synthase IIa differentiate amylopectin structure and starch quality between japonica and indica rice varieties. Plant Mol Biol. 2005;58:213-27 pubmed
    ..are essential not only for the optimal SSIIa activity, but also for the capacity to synthesize indica-type amylopectin. Surprisingly, however, a combination of Phe-781 and Gly-604 could restore about 44% of the SSIIa activity ..
  38. Nakamura Y, Utsumi Y, Sawada T, Aihara S, Utsumi C, Yoshida M, et al. Characterization of the reactions of starch branching enzymes from rice endosperm. Plant Cell Physiol. 2010;51:776-94 pubmed publisher
    ..parameters of all the three starch branching enzyme (BE) isozymes, BEI, BEIIa and BEIIb, from rice with both amylopectin and synthetic amylose as glucan substrate...
  39. Takeda Y, Shibahara S, Hanashiro I. Examination of the structure of amylopectin molecules by fluorescent labeling. Carbohydr Res. 2003;338:471-5 pubmed
    b>Amylopectin molecules from rice, maize, sweet potato and potato were examined by fluorescent labeling followed by gel-permeation HPLC. The number-average degree of polymerization (dp(n)) was determined to be in range of 9600-15,900...
  40. Kamasaka H, Sugimoto K, Takata H, Nishimura T, Kuriki T. Bacillus stearothermophilus neopullulanase selective hydrolysis of amylose to maltose in the presence of amylopectin. Appl Environ Microbiol. 2002;68:1658-64 pubmed
    The specificity of Bacillus stearothermophilus TRS40 neopullulanase toward amylose and amylopectin was analyzed...
  41. Nakamura Y. Towards a better understanding of the metabolic system for amylopectin biosynthesis in plants: rice endosperm as a model tissue. Plant Cell Physiol. 2002;43:718-25 pubmed
    Starch is made up of amylose (linear alpha-1,4-polyglucans) and amylopectin (alpha-1,6-branched polyglucans)...
  42. Dauvillee D, Colleoni C, Mouille G, Morell M, D Hulst C, Wattebled F, et al. Biochemical characterization of wild-type and mutant isoamylases of Chlamydomonas reinhardtii supports a function of the multimeric enzyme organization in amylopectin maturation. Plant Physiol. 2001;125:1723-31 pubmed
    ..the mutant enzyme was demonstrated to digest phytoglycogen to completion in vitro, we propose that its inability to do so in vivo supports a function of the enzyme complex architecture in the processing of pre-amylopectin chains.
  43. Ryan S, Fitzgerald G, van Sinderen D. Screening for and identification of starch-, amylopectin-, and pullulan-degrading activities in bifidobacterial strains. Appl Environ Microbiol. 2006;72:5289-96 pubmed
    ..screened for alpha-amylase and/or pullulanase activity by investigating their capacities to utilize starch, amylopectin, or pullulan. Of the 42 bifidobacterial strains tested, 19 were capable of degrading potato starch...
  44. Wattebled F, Dong Y, Dumez S, Delvalle D, Planchot V, Berbezy P, et al. Mutants of Arabidopsis lacking a chloroplastic isoamylase accumulate phytoglycogen and an abnormal form of amylopectin. Plant Physiol. 2005;138:184-95 pubmed
    ..In addition, the residual amylopectin structure in the corresponding mutant lines displays a strong modification when compared to the wild type, ..
  45. Critchley J, Zeeman S, Takaha T, Smith A, Smith S. A critical role for disproportionating enzyme in starch breakdown is revealed by a knock-out mutation in Arabidopsis. Plant J. 2001;26:89-100 pubmed
    ..During the diurnal cycle, the amount of leaf starch is higher in dpe1-1 than in wild type and the amylose to amylopectin ratio is increased, but amylopectin structure is unaltered...
  46. Damager I, Denyer K, Motawia M, Møller B, Blennow A. The action of starch synthase II on 6"'-alpha-maltotriosyl-maltohexaose comprising the branch point of amylopectin. Eur J Biochem. 2001;268:4878-84 pubmed
    ..Compared to the surface exposed alpha-glucan chains of the granule bound amylopectin molecules, all three soluble oligosaccharides tested were poor primers for SSII...
  47. Rydberg U, Andersson L, Andersson R, Aman P, Larsson H. Comparison of starch branching enzyme I and II from potato. Eur J Biochem. 2001;268:6140-5 pubmed
    ..that SBE I was more active than SBE II on an amylose substrate, whereas SBE II was more active than SBE I on an amylopectin substrate. Both enzymes were stimulated by the presence of phosphate...
  48. Wang X, Conway P, Brown I, Evans A. In vitro utilization of amylopectin and high-amylose maize (Amylomaize) starch granules by human colonic bacteria. Appl Environ Microbiol. 1999;65:4848-54 pubmed
    ..types of human colonic bacteria were screened for their capacity to utilize soluble starch, gelatinized amylopectin maize starch, and high-amylose maize starch granules by measuring the clear zones on starch agar plates...
  49. Streb S, Delatte T, Umhang M, Eicke S, Schorderet M, Reinhardt D, et al. Starch granule biosynthesis in Arabidopsis is abolished by removal of all debranching enzymes but restored by the subsequent removal of an endoamylase. Plant Cell. 2008;20:3448-66 pubmed publisher
    Several studies have suggested that debranching enzymes (DBEs) are involved in the biosynthesis of amylopectin, the major constituent of starch granules...
  50. Commuri P, Keeling P. Chain-length specificities of maize starch synthase I enzyme: studies of glucan affinity and catalytic properties. Plant J. 2001;25:475-86 pubmed
    ..Greater affinity was displayed for the amylopectin fraction of starch as compared to amylose, whereas glycogen revealed the lowest affinity...
  51. Wu A, Gilbert R. Molecular weight distributions of starch branches reveal genetic constraints on biosynthesis. Biomacromolecules. 2010;11:3539-47 pubmed publisher
    ..The model implies that all native-starch amylopectin CLDs are confined to a line in this phase diagram, an inference supported by fitting data for a wide range of ..
  52. Vermeylen R, Goderis B, Reynaers H, Delcour J. Amylopectin molecular structure reflected in macromolecular organization of granular starch. Biomacromolecules. 2004;5:1775-86 pubmed
    ..amorphous lamellae, envisaged by the lamellar helical model, explains the relative acid resistance of linear amylopectin chains with DP > 20, observed in lintners of B-type starches...
  53. Wu A, Morell M, Gilbert R. A parameterized model of amylopectin synthesis provides key insights into the synthesis of granular starch. PLoS ONE. 2013;8:e65768 pubmed publisher
    ..The chain-length distribution (CLD) of amylopectin in cereal endosperm is modeled here on the basis that the CLD is produced by concerted actions of three enzyme ..