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REVIEW

Progress in comprehending the phytate–phytase axis in chicken-meat production

Amy F. Moss A , Sonia Yun Liu A and Peter H. Selle A B
+ Author Affiliations
- Author Affiliations

A Poultry Research Foundation, within The University of Sydney, 425 Werombi Road, Camden, NSW 2570, Australia.

B Corresponding author. Email: peter.selle@sydney.edu.au

Animal Production Science 58(10) 1767-1778 https://doi.org/10.1071/AN17594
Submitted: 31 August 2017  Accepted: 7 December 2017   Published: 21 February 2018

Abstract

After an extended delay, the level of acceptance of exogenous phytases by the global chicken-meat industry is now almost complete. Contemporary bacterial phytases degrade phytate primarily in the gizzard. The extent of phytate degradation determines the extent to which phytate-bound phosphorus (P) is liberated; however, studies designed to investigate phytate degradation along the digestive tract have generated some confusing outcomes. This may be related to the reactivity of the phytate moiety, coupled with problems with inert dietary markers and perhaps a lack of complete and uniform extractions of phytate from digesta due to variations in digesta pH and phytate solubility. Quite recently, phytase was shown to have profound impacts on sodium (Na) digestibility coefficients in four segments of the small intestine. This has obvious implications for intestinal uptakes of glucose and amino acids via their respective Na+-dependent transport systems and it is possible that phytate and phytase have reciprocal impacts on ‘sodium pump’ (Na+, K+-ATPase) activity. It has been recently demonstrated unequivocally that phytase has the capacity to increase amino acid digestibility coefficients to the extent that phytase may generate a ‘proximal shift’ in the sites of amino acid absorption. The impact of phytase on starch digestibility is more equivocal and phytase responses may stem more from enhanced glucose absorption rather than starch digestion. The acceptance of phytase is hardly surprising, given its capacity to increase P utilisation coupled with numerous other positive influences that are still being properly realised.

Additional keywords: amino acids, glucose, protein, sodium, starch.


References

Abdollahi MR, Amerah AM, Ravindran V (2016) Influence of whole wheat inclusion and exogenous enzyme supplementation on growth performance and nutrient utilisation in broiler starters. Proceedings, Australian Poultry Science Symposium 27, 223–225.

Abelson PH (1999) A potential phosphate crisis. Science 283, 2015
A potential phosphate crisis.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1MXit12isbg%3D&md5=b40f411a522cc4c89f65e04ab077210fCAS |

Adeola O, Sands JS (2006) Does supplementary microbial phytase improve amino acid utilization? A perspective that it does not. Journal of Animal Science 81, E78–E85.

Allen A, Flemstrom G (2005) Gastroduodenal mucus bicarbonate barrier: protection against acid and pepsin. American Journal of Physiology. Cell Physiology 288, C1–C19.
Gastroduodenal mucus bicarbonate barrier: protection against acid and pepsin.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2MXhtVGjtrY%3D&md5=4d96b89b971a7015184cd8add6e27cc6CAS |

Amerah AM, Plumstead PW, Barnard LP, Kumar A (2014) Effect of calcium level and phytase addition on ileal phytate degradation and amino acid digestibility of broilers fed corn-based diets. Poultry Science 93, 906–915.
Effect of calcium level and phytase addition on ileal phytate degradation and amino acid digestibility of broilers fed corn-based diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2sXhsFChsLzF&md5=b5578fad9ed2aa4c013ff7ccd34a2150CAS |

Angel R, Saylor SL, Vieira SL, Ward N (2011) Effects of a monocomponent protease on performance and protein utilization in 7- to 22-day-old broiler chickens. Poultry Science 90, 2281–2286.
Effects of a monocomponent protease on performance and protein utilization in 7- to 22-day-old broiler chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3MXhtlGrsb%2FK&md5=31f51c7e485318c2d45aceee58a9c2c9CAS |

Antipatis C, Knap I, Pontoppidan K, Valientes RA, Angel R (2013) Exogenous proteases and their interaction with dietary ingredients. Proceedings, Australian Poultry Science Symposium 24, 31–40.

Bae Y, Kim JH, Lee HG (2013) Combined effect of protease and phytase on the solubility of modified soy protein. Journal of Food Biochemistry 37, 511–519.

Baker DH (1998) Beyond phosphorus: phytase effect on protein, energy and trace mineral utilization of swine and poultry. In ‘BASF technical symposium preceding North Carolina swine nutrition conference’, Durham NC. pp. 48–62. (BASF Corporation: Mount Olive, NJ)

Baldwin PM (2001) Starch granule-associated proteins and polypeptides: a review. Stärke 53, 475–503.
Starch granule-associated proteins and polypeptides: a review.Crossref | GoogleScholarGoogle Scholar |

Bedford MR, Cowieson AJ (2010) Additivity of response to different exogenous enzymes in pigs and poultry. In ‘Proceedings, 31st western nutrition conference – challenging conventional nutrition dogma’, Saskatoon, SK. pp. 243–249. (Animal Nutrition Association of Canada: Ottawa, ON)

Bohak Z (1969) Purification and characterization of chicken pepsinogen and chicken pepsin. The Journal of Biological Chemistry 244, 4638–4648.

Bröer S (2008) Amino acid transport across mammalian intestinal and renal epithelia. Physiological Reviews 88, 249–286.
Amino acid transport across mammalian intestinal and renal epithelia.Crossref | GoogleScholarGoogle Scholar |

Bye JW, Cowieson NP, Cowieson AJ, Selle PH, Falconer RJ (2013) Dual effects of sodium phytate on the structural stability and solubility of proteins. Journal of Agricultural and Food Chemistry 61, 290–295.
Dual effects of sodium phytate on the structural stability and solubility of proteins.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XhvVKmsbzE&md5=0d235a25310bd589032ff97f0e4fce55CAS |

Camden BJ, Morel PCH, Thomas DV, Ravindran V, Bedford MR (2001) Effectiveness of exogenous microbial phytase in improving the bioavailabilities of phosphorus and other nutrients in maize–soya-bean meal diets for broilers. Animal Science Journal 73, 289–297.
Effectiveness of exogenous microbial phytase in improving the bioavailabilities of phosphorus and other nutrients in maize–soya-bean meal diets for broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD38XitFChuw%3D%3D&md5=af1ebebf48a4b830094b125e61b557fbCAS |

Camus MC, Laporte JC (1976) Inhibition de la protéolyse pesique par le blé. Rôle de l’acide phytique des issues. Annales de Biologie Animale, Biochimie, Biophysique 16, 719–729.
Inhibition de la protéolyse pesique par le blé. Rôle de l’acide phytique des issues.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE2sXkt1ShtA%3D%3D&md5=aa0a152d3fb90eb1218fb5b4c37ca77bCAS |

Case RM, Harper AA, Scratcherd T (1969) The secretion of electrolytes and enzymes by the pancreas of the anaesthetized cat. The Journal of Physiology 201, 335–348.
The secretion of electrolytes and enzymes by the pancreas of the anaesthetized cat.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaF1MXpsVKiug%3D%3D&md5=324240998477a6a68b2191923afe90c8CAS |

Case RM, Scratcherd T, Wynne RDA (1970) The origin and secretion of pancreatic juice bicarbonate. The Journal of Physiology 210, 1–15.
The origin and secretion of pancreatic juice bicarbonate.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE3cXkvFKnsL4%3D&md5=aa350e95249ed12027ad1000691676b5CAS |

Chen L, Biguang T, Dong H (2016) Regulation of intestinal glucose absorption by ion channels and transporters. Nutrients 43,
Regulation of intestinal glucose absorption by ion channels and transporters.Crossref | GoogleScholarGoogle Scholar |

Cosgrove DJ (1966) The chemistry and biochemistry of inositol polyphosphates. Reviews of Pure and Applied Chemistry 16, 209–224.

Cowieson AJ (2010) Strategic selection of exogenous enzymes for corn/soy-based poultry diets. Journal of Poultry Science 47, 1–7.
Strategic selection of exogenous enzymes for corn/soy-based poultry diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3cXis1Wqu74%3D&md5=c1df7fa8d1e1d93a0131bfd1fe7e93f7CAS |

Cowieson AJ, Ravindran V (2008) Influence of dietary phytic acid and source of microbial phytase on ileal endogenous amino acid flows in broiler chickens. British Journal of Nutrition 87, 2287–2299.

Cowieson AJ, Acamovic T, Bedford MR (2004) The effects of phytase and phytic acid on the loss of endogenous amino acids and minerals from broiler chickens. British Poultry Science 45, 101–108.
The effects of phytase and phytic acid on the loss of endogenous amino acids and minerals from broiler chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXhs1Wgu7c%3D&md5=b2428f4e0ab2acd24720dc973de546d7CAS |

Cowieson AJ, Ravindran V, Selle PH (2008) Influence of dietary phytic acid and source of microbial phytase on ileal endogenous amino acid flows in broiler chickens. Poultry Science 87, 2287–2299.

Cowieson AJ, Bedford MR, Selle PH, Ravindran V (2009) Phytate and microbial phytase: implications for endogenous nitrogen losses and nutrient availability. World’s Poultry Science Journal 65, 401–417.
Phytate and microbial phytase: implications for endogenous nitrogen losses and nutrient availability.Crossref | GoogleScholarGoogle Scholar |

Cowieson AJ, Bedford MR, Ravindran V, Selle PH (2011) Increased dietary sodium chloride concentrations reduce endogenous amino acid flow and influence the physiological response to the ingestion of phytic acid by broiler chickens. British Poultry Science 52, 613–624.
Increased dietary sodium chloride concentrations reduce endogenous amino acid flow and influence the physiological response to the ingestion of phytic acid by broiler chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3MXhtlyhu7nL&md5=5daa04725c9bfed9f578783a855494c8CAS |

Cowieson AJ, Ptak A, Mackowiak P, Sessek M, Pruszynska-Oszmalek E, Zyla K, Swiatkiewicz S, Kaczmarek S, Jozefiak D (2013) The effect of microbial phytase and myo-inositol on performance and blood biochemistry of broiler chickens fed wheat/corn-based diets. Poultry Science 92, 2124–2134.
The effect of microbial phytase and myo-inositol on performance and blood biochemistry of broiler chickens fed wheat/corn-based diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXht1Ois7nK&md5=fc5a04596f701ae50a28cbde98925e35CAS |

Cowieson AJ, Ruckebusch JP, Sorbara JOB, Wilson JW, Guggenbuhl P, Roos FF (2017) A systematic view on the effect of phytase on ileal amino aciddigestibility in broilers. Animal Feed Science and Technology 225, 182–194.
A systematic view on the effect of phytase on ileal amino aciddigestibility in broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2sXisVCnt7o%3D&md5=294e46f28c449fd1d65b01e3467732a0CAS |

Csonka FA, Murphy JC, Jones DB (1926) The iso-electric points of various proteins. The Journal of Biological Chemistry 62, 763–768.

Darby SJ, Platts L, Daniel MS, Cowieson AJ, Falconer RJ (2016) An isothermal titration calorimetry study of phytate binding to lysozyme. Journal of Thermal Analysis and Calorimetry
An isothermal titration calorimetry study of phytate binding to lysozyme.Crossref | GoogleScholarGoogle Scholar |

de Boland AR, Garner GB, O’Dell BL (1975) Identification and properties of ‘phytate’ in cereal grains and oilseed products. Journal of Agricultural and Food Chemistry 23, 1186–1189.
Identification and properties of ‘phytate’ in cereal grains and oilseed products.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE2MXlvFWmtbg%3D&md5=2571aaa39444e5ee1a5da76627add659CAS |

de Vries S, Kwakkel RP, Pustjens AM, Kabel MA, Hendriks WH, Gerrits WJJ (2014) Separation of digesta fractions complicates estimation of ileal digestibility using marker methods with Cr2O3 and cobalt–ethylenediamine tetraacetic acid in broiler chickens. Poultry Science 93, 2010–2017.
Separation of digesta fractions complicates estimation of ileal digestibility using marker methods with Cr2O3 and cobalt–ethylenediamine tetraacetic acid in broiler chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXhtlaks7nM&md5=5ac28e9c036da90be17c281b4320e468CAS |

Demjen AP, Thompson LU (1991) Calcium and phytic acid independently lower the glycemic response to a glucose load. In ‘Proceedings, 34th Canadian Federation of Biological Sciences’. 34, 53 (Abstr.).

Deshpande SS, Cheryan M (1984) Effects of phytic acid, divalent cations and their interactions on a-amylase activity. Journal of Food Science 49, 516–519.
Effects of phytic acid, divalent cations and their interactions on a-amylase activity.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL2cXhvVOqsLg%3D&md5=a0d911f171e26e6e646618db3aa1e449CAS |

Dilworth LL, Omoruyi FO, Asemota HN (2005) Digestive and absorptive enzymes in rats fed phytic acid extract from sweet potato (Ipomoea batatas). Diabetologia Croatica 34, 59–65.

Engberg RM, Hedemann MS, Steenfeldt S, Jensen BB (2004) Influence of whole wheat and xylanase on broiler performance and microbial composition and activity in the digestive tract. Poultry Science 83, 925–938.
Influence of whole wheat and xylanase on broiler performance and microbial composition and activity in the digestive tract.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXltlCktbo%3D&md5=cdb9396c5fdbbe8a217660a3a70355b6CAS |

Farhadi D, Karimi A, Sadeghi G, Rostamzadeh J, Bedford MR (2017) Effects of a high dose of microbial phytase and myo-inositol supplementation on growth performance, tibia mineralization, nutrient digestibility, litter moisture content, and foot problems in broiler chickens fed phosphorus-deficient diets. Poultry Science
Effects of a high dose of microbial phytase and myo-inositol supplementation on growth performance, tibia mineralization, nutrient digestibility, litter moisture content, and foot problems in broiler chickens fed phosphorus-deficient diets.Crossref | GoogleScholarGoogle Scholar |

Farrell DJ, Martin E, du Preez JJ, Bongarts M, Betts M, Sudaman A, Thomson E (1993) The beneficial effects of a microbial phytase in diets of broiler chickens and ducklings. Journal of Animal Physiology and Animal Nutrition 69, 278–283.
The beneficial effects of a microbial phytase in diets of broiler chickens and ducklings.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2cXlsFKgsb4%3D&md5=54b4ec103ef6e0018ccb9ec91c6c624cCAS |

Gal-Garber O, Mabjeesh SJ, Sklan D, Uni Z (2003) Nutrient transport in the small intestine: Na+, K+-ATPase expression and activity in the small intestine of the chicken as influenced by dietary sodium. Poultry Science 82, 1127–1133.
Nutrient transport in the small intestine: Na+, K+-ATPase expression and activity in the small intestine of the chicken as influenced by dietary sodium.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3sXmsVWkt7k%3D&md5=3ab544d05910318a22cd11124d1dab8bCAS |

Gehring CK, Bedford MR, Dozier WA (2013) Interactive effects of phytase and xylanase supplementation with extractable salt-soluble protein content of corn in diets with adequate calcium and nonphytate phosphorus fed to broilers. Poultry Science 92, 1858–1869.
Interactive effects of phytase and xylanase supplementation with extractable salt-soluble protein content of corn in diets with adequate calcium and nonphytate phosphorus fed to broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXhtVGlsLjJ&md5=dabba982d61bbf370410eca4417ff969CAS |

Glynn IM (1993) All hands to the sodium pump. The Journal of Physiology 462, 1–30.
All hands to the sodium pump.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK3szitlSjtg%3D%3D&md5=7747c9dbf31e4ebf30fc16281bb0f127CAS |

Grynspan F, Cheryan M (1983) Calcium phytate: effect of pH and molar ratio on in vitro solubility. Journal of the American Oil Chemists’ Society 60, 1761–1764.
Calcium phytate: effect of pH and molar ratio on in vitro solubility.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL3sXlvFehu7s%3D&md5=28e6af280fa77466128ac1927ed510e7CAS |

Kaufman HW, Kleinberg I (1971) Effect of pH on calcium binding by phytic acid and its inositol phosphoric acid derivatives and on the solubility of their calcium salts. Archives of Oral Biology 16, 445–460.
Effect of pH on calcium binding by phytic acid and its inositol phosphoric acid derivatives and on the solubility of their calcium salts.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE3MXktFGhs78%3D&md5=f49bfb813a1de05c7ae5c3230920010bCAS |

Kiarie E, Romero LF, Ravindran V (2014) Growth performance, nutrient utilization, and digesta characteristics in broiler chickens fed corn or wheat diets without or with supplemental xylanase. Poultry Science 93, 1186–1196.
Growth performance, nutrient utilization, and digesta characteristics in broiler chickens fed corn or wheat diets without or with supplemental xylanase.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXosFCgu7c%3D&md5=72dac71ab5aac245b1252722fbf0f6d4CAS |

Knuckles BE, Kuzmicky DD, Gumbmann MR, Betschart AA (1989) Effect of myo-inositol phosphate esters on in vitro and in vivo digestion of protein. Journal of Food Science 54, 1348–1350.
Effect of myo-inositol phosphate esters on in vitro and in vivo digestion of protein.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL1MXmtlSrt7w%3D&md5=807a6e98399f194cf19e42336c4e366fCAS |

Krehbiel CR, Matthews JC (2003) Absorption of amino acids and peptides. In ‘Amino acids in animal nutrition’. 2nd edn. (Ed. JPF D’Mello) pp. 41–70. (CAB International: Wallingford, UK)

Lee SA, Bedford MR (2016) Inositol: an effective growth promotor? World’s Poultry Science Journal 72, 743–759.
Inositol: an effective growth promotor?Crossref | GoogleScholarGoogle Scholar |

Li W, Angel R, Kim SW, Jimenez-Moreno E, Proszkowiec-Weglarz M, Plumstead PW (2015) Age and adaptation to Ca and P deficiencies: 2. Impacts on amino acid digestibility and phytase efficacy in broilers. Poultry Science 94, 2917–2931.
Age and adaptation to Ca and P deficiencies: 2. Impacts on amino acid digestibility and phytase efficacy in broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2sXhvFCgsLnI&md5=7943882692cc7c269ae380b7c86cc234CAS |

Lien KA, Sauer WC, Fenton M (1997) Mucin output in ileal digesta of pigs fed a protein-free diet. Zeitschrift fur Ernahrungswissenschaft 36, 182–190.
Mucin output in ileal digesta of pigs fed a protein-free diet.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2sXlt1ahsro%3D&md5=eaa201ab77c6fea6f699ca9faf668594CAS |

Liu SY, Selle PH (2015) A consideration of starch and protein digestive dynamics in chicken-meat production. World’s Poultry Science Journal 71, 297–310.
A consideration of starch and protein digestive dynamics in chicken-meat production.Crossref | GoogleScholarGoogle Scholar |

Liu N, Ru YJ, Li FD, Cowieson AJ (2008) Effect of diet containing phytate and phytase on the activity and messenger ribonucleic acid expression of carbohydrase and transporter in chickens. Journal of Animal Science 86, 3432–3439.
Effect of diet containing phytate and phytase on the activity and messenger ribonucleic acid expression of carbohydrase and transporter in chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1cXhsV2jsb3P&md5=18264d9b354ab8fcdac64c8bcfbed904CAS |

Liu SY, Cadogan DJ, Peron A, Truong HH, Selle PH (2014) Effects of phytase supplementation on growth performance, nutrient utilization and digestive dynamics of starch and protein in broiler chickens offered maize-, sorghum- and wheat-based diets. Animal Feed Science and Technology 197, 164–175.
Effects of phytase supplementation on growth performance, nutrient utilization and digestive dynamics of starch and protein in broiler chickens offered maize-, sorghum- and wheat-based diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXhsV2nsb%2FL&md5=46484c9e8cdbbc599796756b5b55c126CAS |

Liu SY, Truong HH, Selle PH (2015) Whole grain feeding for chicken-meat production: possible mechanisms driving enhanced energy utilisation and feed conversion. Animal Production Science 55, 559–572.
Whole grain feeding for chicken-meat production: possible mechanisms driving enhanced energy utilisation and feed conversion.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2MXlsV2jtr4%3D&md5=0c30915c0bf4b213d15664d6543252dbCAS |

Liu SY, Cowieson AJ, Selle PH (2016) The influence of meat-and-bone meal and exogenous phytase on growth performance, bone mineralisation and digestibility coefficients of protein (N), amino acids and starch in broiler chickens. Animal Nutrition 2, 86–92.
The influence of meat-and-bone meal and exogenous phytase on growth performance, bone mineralisation and digestibility coefficients of protein (N), amino acids and starch in broiler chickens.Crossref | GoogleScholarGoogle Scholar |

Lonnerdal B, Sandberg A-S, Sandstrom B, Kunz C (1989) Inhibitory effects of phytic acid and other inositol phosphates on zinc and calcium absorption in suckling rats. The Journal of Nutrition 119, 211–214.

Luttrell BM (1993) The biological relevance of the binding of calcium ions by inositol phosphates. The Journal of Biological Chemistry 268, 1521–1524.

Maenz DD, Classen HL (1998) Phytase activity in the small intestinal brush border membrane of the chicken. Poultry Science 77, 557–563.
Phytase activity in the small intestinal brush border membrane of the chicken.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1cXks1aqt78%3D&md5=50cad448d2ac34e414a2138e28322730CAS |

Marini MA, Evans WJ, Morris NM (1985) Calometric and potentiometric studies on the binding of calcium by phytic acid. Journal of Applied Biochemistry 7, 180–191.

Martinez-Amezcua C, Parsons CM, Baker DH (2006) Effect of microbial phytase on phosphorus bioavailability, apparent metabolizable energy, and amino acid digestibility in distillers dried grains with solubles in chicks. Poultry Science 85, 470–475.
Effect of microbial phytase on phosphorus bioavailability, apparent metabolizable energy, and amino acid digestibility in distillers dried grains with solubles in chicks.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28Xis1aqu7c%3D&md5=792d9da404816d846fc8d5a275756d47CAS |

Menezes-Blackburn D, Gabler S, Greiner R (2015) Performance of seven commercial phytases in an in vitro simulation of poultry digestive tract. Journal of Agricultural and Food Chemistry 63, 6142–6149.
Performance of seven commercial phytases in an in vitro simulation of poultry digestive tract.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2MXhtVOit73I&md5=4588057282384e72c0a63eca389b047cCAS |

Morgan EL, Mace OJ, Affleck J, Kellett GL (2007) Apical GLUT2 and Cav1.3: regulation of rat intestinal glucose and calcium absorption. The Journal of Physiology 580, 593–604.
Apical GLUT2 and Cav1.3: regulation of rat intestinal glucose and calcium absorption.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXltFeitbY%3D&md5=be3c0e35654fbb603c2b682aafea8a0bCAS |

Moss AF, Chrystal PV, Truong HH, Liu SY, Selle PH (2017a) Effects of pre- and post-pelleting whole grain additions and exogenous phytase on the performance of broiler chickens offered wheat-based diets plus phytase and protease supplementation of pre-pelleted whole grain diets. Animal Feed Science and Technology 234, 139–150.
Effects of pre- and post-pelleting whole grain additions and exogenous phytase on the performance of broiler chickens offered wheat-based diets plus phytase and protease supplementation of pre-pelleted whole grain diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2sXhsFOnsL7M&md5=42c8f7b86a0ef428b7dcc080c60e4f8aCAS |

Moss AF, Sydenham CJ, Truong HH, Liu SY, Selle PH (2017b) The interactions of exogenous phytase with whole grain feeding and effects of barley as the whole grain component in broiler diets based on wheat, sorghum and wheat-sorghum blends. Animal Feed Science and Technology 227, 1–12.
The interactions of exogenous phytase with whole grain feeding and effects of barley as the whole grain component in broiler diets based on wheat, sorghum and wheat-sorghum blends.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2sXkslSgsr0%3D&md5=5c56f01e94ff845ea421d082b9085e38CAS |

Nelson TS (1984) Available calcium for poultry. In ‘Proceedings, Florida nutrition conference for feed manufacturers’. pp. 1–7.

Onyango EM, Asem EK, Adeola O (2009) Phytic acid increases mucin and endogenous amino acid losses from the gastrointestinal tract of chickens. British Journal of Nutrition 101, 836–842.
Phytic acid increases mucin and endogenous amino acid losses from the gastrointestinal tract of chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXktleisr8%3D&md5=00566618a3c1e0fd98b5c0d1d8562713CAS |

Rajendran S, Prakash V (1993) Kinetics and thermodynamics of the mechanism of interaction of sodium phytate with a-globulin. Biochemistry 32, 3474–3478.
Kinetics and thermodynamics of the mechanism of interaction of sodium phytate with a-globulin.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3sXhvVKitrY%3D&md5=0724eae505a770fc837be99100b7b5e5CAS |

Ravindran V (1995) Phytases in poultry nutrition. An overview. Proceedings, Australian Poultry Science Symposium 7, 135–139.

Ravindran V, Selle PH, Bryden WL (1999) Effects of phytase supplementation, individually and in combination, with glycanase on the nutritive value of wheat and barley. Poultry Science 78, 1588–1595.
Effects of phytase supplementation, individually and in combination, with glycanase on the nutritive value of wheat and barley.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1MXnsFSls7k%3D&md5=3c31bddf1b071557b170c31c08c00b00CAS |

Ravindran V, Cabahug S, Ravindran G, Selle PH, Bryden WL (2000) Response of broiler chickens to microbial phytase supplementation as influenced by dietary phytic acid and non-phytate phosphorus levels. II. Effects on apparent metabolisable energy, nutrient digestibility and nutrient retention. British Poultry Science 41, 193–200.
Response of broiler chickens to microbial phytase supplementation as influenced by dietary phytic acid and non-phytate phosphorus levels. II. Effects on apparent metabolisable energy, nutrient digestibility and nutrient retention.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXksVCmtLc%3D&md5=757e6a4dfdbf54f092aed379105f684bCAS |

Ravindran V, Morel PCH, Partridge GG, Hruby M, Sands JS (2006) Influence of an E. coli-derived phytase on nutrient utilization in broiler starters fed diets containing varying concentrations of phytic acid. Poultry Science 85, 82–89.
Influence of an E. coli-derived phytase on nutrient utilization in broiler starters fed diets containing varying concentrations of phytic acid.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28XltVKrsA%3D%3D&md5=b10424e94cb5e2668f6f056a61491eb3CAS |

Ravindran V, Cowieson AJ, Selle PH (2008) Influence of dietary electrolyte balance and microbial phytase on growth performance, nutrient utilization and excreta quality of broiler chickens. Poultry Science 87, 677–688.

Rickard SE, Thompson LU (1997) Interactions and biological effects of phytic acid. In ‘Antinutrients and phytochemicals in food’. (Ed. F Shahidi) pp. 294–312. (American Chemical Society: Washington, DC)

Romero LF, Plumstead PW (2013) Bio-efficacy of feed proteases and their interaction with other feed enzymes. Proceedings, Australian Poultry Science Symposium 24, 23–30.

Schwarz G, Schöner F-J, Weiske H, Hoppe (1994) Evaluation of a possible interaction between microbial phytase and two carbohydrase preparations in broilers. Proceedings, Society of Nutrition Physiology 2, 47

Sebastian A, Touchburn SP, Chavez ER, Lague PC (1997) Apparent digestibility of protein and amino acids in broiler chickens fed a corn-soybean diet supplemented with microbial phytase. Poultry Science 76, 1760–1769.
Apparent digestibility of protein and amino acids in broiler chickens fed a corn-soybean diet supplemented with microbial phytase.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1cXjvFKgsrY%3D&md5=67013dfcc01bb6cdf9fafb95d1da10e0CAS |

Selle PH, Ravindran V (2007) Microbial phytase in poultry nutrition. Animal Feed Science and Technology 135, 1–41.
Microbial phytase in poultry nutrition.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXkt1artb8%3D&md5=f72fc32508501d3d4105ce75c31ab388CAS |

Selle PH, Ravindran V, Caldwell RA, Bryden WL (2000) Phytate and phytase: consequences for protein utilisation. Nutrition Research Reviews 13, 255–278.
Phytate and phytase: consequences for protein utilisation.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3MXhtlKmtr8%3D&md5=16537b90b64aac4d5c87b0187bbfa769CAS |

Selle PH, Partridge GG, Ravindran V (2009) Beneficial effects of xylanase and/or phytase inclusions on ileal amino acid digestibility energy utilisation mineral retention and growth performance in wheat-based broiler diets. Animal Feed Science and Technology 153, 303–313.
Beneficial effects of xylanase and/or phytase inclusions on ileal amino acid digestibility energy utilisation mineral retention and growth performance in wheat-based broiler diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD1MXhtV2ku77K&md5=93bdb5aad87cf9453a29ace631f730f8CAS |

Selle PH, Cowieson AJ, Cowieson NP, Ravindran V (2012) Protein–phytate interactions in pig and poultry nutrition: a reappraisal. Nutrition Research Reviews 25, 1–17.
Protein–phytate interactions in pig and poultry nutrition: a reappraisal.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XpvVOisb0%3D&md5=ac6e1d786131dafc2721623727717f2eCAS |

Shafey TM, McDonald MW, Dingle JG (1991) Effects of dietary calcium and available phosphorus concentration on digesta pH and on the availability of calcium, iron, magnesium, and zinc from the intestinal contents of meat chickens. British Poultry Science 32, 185–194.
Effects of dietary calcium and available phosphorus concentration on digesta pH and on the availability of calcium, iron, magnesium, and zinc from the intestinal contents of meat chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3MXmslymurc%3D&md5=dfba1a22e5a6e0390f0070af89375e75CAS |

Sharma CB, Goel M, Irshad M (1978) Myoinositol hexaphosphate as a potential inhibitor of α-amylases. Phytochemistry 17, 201–204.
Myoinositol hexaphosphate as a potential inhibitor of α-amylases.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE1cXhtFKqsb8%3D&md5=cda62c1a1729e4f05477fc393a83b1faCAS |

Simons PCM, Versteegh HAJ, Jongbloed AW, Kemme PA, Slump P, Bos KD, Wolters MGE, Beudeker RF, Verschoor GJ (1990) Improvement of phosphorus availability by microbial phytase in broilers and pigs. British Journal of Nutrition 64, 525–540.
Improvement of phosphorus availability by microbial phytase in broilers and pigs.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3cXlslSrsrc%3D&md5=ff5b3b0a6505b9e6d4998eda4ca03213CAS |

Singh Y, Amerah AM, Ravindran V (2014) Whole grain feeding: methodologies and effects on performance, digestive tract development and nutrient utilisation of poultry. Animal Feed Science and Technology 190, 1–18.
Whole grain feeding: methodologies and effects on performance, digestive tract development and nutrient utilisation of poultry.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXitV2ltL4%3D&md5=4a745ff9acfc7e38036a3bed87092c3aCAS |

Stefanello C, Vieira SL, Santiago GO, Kindelin L, Sorbara JOB, Cowieson AJ (2015) Starch digestibility, energy utilisation, and growth performance of broilers feds corn–soybean based diets supplemented with enzymes. Poultry Science 94, 2472–2479.
Starch digestibility, energy utilisation, and growth performance of broilers feds corn–soybean based diets supplemented with enzymes.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC28XhvFOrtbjL&md5=d106cda2717900a654adf7d31a0ada8cCAS |

Takemasa M, Murakami H, Yamazaki M (1996) Reduction of phosphorus excretion in chicks by the addition of yeast phytase. Japanese Poultry Science 33, 104–111.
Reduction of phosphorus excretion in chicks by the addition of yeast phytase.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK28XisVGlurc%3D&md5=38051677835f72c4b75177b047a314afCAS |

Tamim NM, Angel R, Christman M (2004) Influence of dietary calcium and phytase on phytate phosphorus hydrolysis in broiler chickens. Poultry Science 83, 1358–1367.
Influence of dietary calcium and phytase on phytate phosphorus hydrolysis in broiler chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXntFGlsro%3D&md5=61956520e42cd30ab11a6c92b25c6423CAS |

Therien AG, Blostein R (2000) Mechanisms of sodium pump regulation. The American Journal of Physiology 279, C541–C566.
Mechanisms of sodium pump regulation.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXmvVGktbc%3D&md5=0203d33d6c7058fa01ada1c351cf0b9eCAS |

Thompson LU (1988a) Antinutrients and blood glucose. Food Technology 42, 123–131.

Thompson LU (1988b) Phytic acid: a factor influencing starch digestibility and blood glucose response. In ‘Phytic acid: chemistry and applications’. (Ed. E Graf) pp. 173–194. (Pilatus Press: Minneapolis, MN)

Thompson LU, Yoon JH (1984) Starch digestibility as affected by polyphenols and phytic acid. Journal of Food Science 49, 1228–1229.
Starch digestibility as affected by polyphenols and phytic acid.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaL2cXlvFSjsbo%3D&md5=1b92bb05799ebc938c7c6a7365ccaf2aCAS |

Thorne MJ, Thompson LU, Jenkins DJA (1983) Factors affecting starch digestibility and the glycemic response with special reference to legumes. The American Journal of Clinical Nutrition 38, 481–488.

Truong HH, Yu S, Peron A, Cadogan DJ, Khoddami A, Roberts TH, Liu SY, Selle PH (2014) Phytase supplementation of maize-, sorghum- and wheat-based broiler diets with identified starch pasting properties influences phytate (IP6) and sodium jejunal and ileal digestibility. Animal Feed Science and Technology 198, 248–256.
Phytase supplementation of maize-, sorghum- and wheat-based broiler diets with identified starch pasting properties influences phytate (IP6) and sodium jejunal and ileal digestibility.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXhvVWmtrvJ&md5=b79aa059434abbd207719be026c5451eCAS |

Truong HH, Bold RM, Liu SY, Selle PH (2015) Standard phytase inclusion in maize-based broiler diets enhances digestibility coefficients of starch, amino acids and sodium in four small intestinal segments and digestive dynamics of starch and protein. Animal Feed Science and Technology 209, 240–248.
Standard phytase inclusion in maize-based broiler diets enhances digestibility coefficients of starch, amino acids and sodium in four small intestinal segments and digestive dynamics of starch and protein.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2MXhsFeitLrJ&md5=598aa6128c2d618c0b6dd6a91f35c0f0CAS |

Truong HH, Cadogan DJ, Liu SY, Selle PH (2016) Addition of sodium metabisulfite and microbial phytase, individually and in combination, to a sorghum-based diet for broiler chickens from 7 to 28 days post-hatch. Animal Production Science 56, 1484–1491.
Addition of sodium metabisulfite and microbial phytase, individually and in combination, to a sorghum-based diet for broiler chickens from 7 to 28 days post-hatch.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC28Xht1Krs7bL&md5=93902cdfb120fccfd1d09e8cc3388797CAS |

Truong HH, Yu S, Moss AF, Partridge GG, Liu SY, Selle PH (2017a) Phytase inclusions of 500 and 2000 FTU/kg in maize-based broiler diets impact on growth performance, nutrient utilisation, digestive dynamics of starch, protein (N), sodium and IP6 phytate degradation in the gizzard and four small intestinal segments. Animal Feed Science and Technology 223, 13–22.
Phytase inclusions of 500 and 2000 FTU/kg in maize-based broiler diets impact on growth performance, nutrient utilisation, digestive dynamics of starch, protein (N), sodium and IP6 phytate degradation in the gizzard and four small intestinal segments.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC28XhvVynt77E&md5=1a1ed7237447cdcade41c48513f3f991CAS |

Truong HH, Moss AF, Liu SY, Selle PH (2017b) Pre- and post-pellet whole grain inclusions enhance feed conversion efficiency, energy utilisation and gut integrity in broiler chickens offered wheat-based diets. Animal Feed Science and Technology 224, 115–123.
Pre- and post-pellet whole grain inclusions enhance feed conversion efficiency, energy utilisation and gut integrity in broiler chickens offered wheat-based diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2sXmsFSgtA%3D%3D&md5=b51c7ee1bb60138cfa7bccb1a613d178CAS |

Vaintraub IA, Bulmaga VP (1991) Effect of phytate on the in vitro activity of digestive proteinases. Journal of Agricultural and Food Chemistry 39, 859–861.
Effect of phytate on the in vitro activity of digestive proteinases.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3MXit1Ols7Y%3D&md5=9835f2620d84f6ef3f4c6529769ec57aCAS |

Viveros A, Centeno C, Brenes A, Canales R, Lozana A (2000) Phytase and phosphatase activities in plant feedstuffs. Journal of Agricultural and Food Chemistry 48, 4009–4013.
Phytase and phosphatase activities in plant feedstuffs.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3cXlt1entrg%3D&md5=cb1cd0806897ea1455797dc31c706773CAS |

Walk CL, Santos TT, Bedford MR (2014) Influence of superdoses of a novel microbial phytase on growth performance, tibia ash, and gizzard phytate and inositol in young broilers. Poultry Science 93, 1172–1177.
Influence of superdoses of a novel microbial phytase on growth performance, tibia ash, and gizzard phytate and inositol in young broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXosFCns70%3D&md5=31f412a6ff1228f158fcbae938d42fdaCAS |

Watford M, Lund P, Krebs KA (1979) Isolation and metabolic characteristics of rat and chicken enterocytes. The Biochemical Journal 178, 589–596.
Isolation and metabolic characteristics of rat and chicken enterocytes.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaE1MXlt1Wjtbg%3D&md5=aad68691010abe79e043fade69af6103CAS |

Wise A (1983) Dietary factors determining the biological activity of phytates. Nutrition Abstracts and Reviews: Clinical Nutrition 53, 791–806.

Wright EM (1993) The intestinal Na+/glucose cotransporter. Annual Review of Physiology 55, 575–589.
The intestinal Na+/glucose cotransporter.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK3sXisFWrt7o%3D&md5=6484dce1247844a7dc08f206c96fd723CAS |

Wu G (1998) Intestinal mucosal amino acid catabolism. The Journal of Nutrition 128, 1249–1252.

Wu YB, Ravindran V (2004) Influence of whole wheat inclusion and xylanase supplementation on the performance digestive tract measurements and carcass characteristics of broiler chickens. Animal Feed Science and Technology 116, 129–139.
Influence of whole wheat inclusion and xylanase supplementation on the performance digestive tract measurements and carcass characteristics of broiler chickens.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXmtVagt7w%3D&md5=23b8b5c584eac48feff98b4745ac1798CAS |

Wu YB, Ravindran V, Thomas DG, Birtles MJ, Hendriks WH (2004) Influence of phytase and xylanase, individually or in combination, on performance, apparent metabolisable energy, digestive tract measurements and gut morphology of broilers. British Poultry Science 45, 385–394.
Influence of phytase and xylanase, individually or in combination, on performance, apparent metabolisable energy, digestive tract measurements and gut morphology of broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2cXlsFyqt70%3D&md5=b7581efc5315d9af8dfbc349c2b878e3CAS |

Yu S, Cowieson A, Gilbert C, Plumsted P, Dalsgaard S (2012) Interactions of phytate and myo-inositol phosphate esters (IP1-5) including IP5 isomers with dietary protein and iron and inhibition of pepsin. Journal of Animal Science 90, 1824–1832.
Interactions of phytate and myo-inositol phosphate esters (IP1-5) including IP5 isomers with dietary protein and iron and inhibition of pepsin.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XpsFWgsbw%3D&md5=a974154acf00f6def3faabd9d0ba0464CAS |

Yu S, Kvidtgaard MF, Isaksen MF, Dalsgaard S (2014) Characterisation of a mutant buttiauxella phytase using phytic acid and phytic acid-protein complex as substrates. Animal Science Letters 1, 18–32.

Zaefarian F, Romero LF, Ravindran V (2013) Influence of a microbial phytase on the performance and the utilisation of energy, crude protein and fatty acids of young broilers fed on phosphorus-adequate maize- and wheat-based diets. British Poultry Science 54, 653–660.
Influence of a microbial phytase on the performance and the utilisation of energy, crude protein and fatty acids of young broilers fed on phosphorus-adequate maize- and wheat-based diets.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXht1CktbzM&md5=f3499447a314d4dc14d45934eedcfc91CAS |

Zeller E, Schollenberger M, Kuhn I, Rodehutscord M (2015) Hydrolysis of phytate and formation of inositol phosphate isomers without or with supplemented phytases in different segments of the digestive tract of broilers. Journal of Nutritional Science 4,
Hydrolysis of phytate and formation of inositol phosphate isomers without or with supplemented phytases in different segments of the digestive tract of broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2MXitlemur0%3D&md5=d04d4cb937ff19dfba225f18bba29eecCAS |

Zyla K, Gogol D, Koreleski J, Swiatkiewicz S, Ledoux DR (1999a) Simultaneous application of phytase and xylanase to broiler feeds based on wheat: in vitro measurements of phosphorus and pentose release from wheats and wheat-based feeds. Journal of the Science of Food and Agriculture 79, 1832–1840.
Simultaneous application of phytase and xylanase to broiler feeds based on wheat: in vitro measurements of phosphorus and pentose release from wheats and wheat-based feeds.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1MXmslent7k%3D&md5=8ff7d0599931be70fff96d28a2d9754aCAS |

Zyla K, Gogol D, Koreleski J, Swiatkiewicz S, Ledoux DR (1999b) Simultaneous application of phytase and xylanase to broiler feeds based on wheat: feeding experiment with growing broilers. Journal of the Science of Food and Agriculture 79, 1841–1848.
Simultaneous application of phytase and xylanase to broiler feeds based on wheat: feeding experiment with growing broilers.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK1MXmslent7Y%3D&md5=93974ceb03dfbdaadceac9a656498acdCAS |