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Review: Phosphorus in Fish Nutrition

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view of the evidence of <strong>in</strong>teraction and balance among food constituents, . . . optimum Ca and P contents <strong>in</strong> diet<br />

vary every time the other elements of diet are changed." Sugiura et al. (1998) calculated the <strong>in</strong>teraction<br />

(<strong>in</strong>hibition) potential of each feed <strong>in</strong>gredient by measur<strong>in</strong>g m<strong>in</strong>eral absorption when a semi-purified bas al diet was<br />

fed alone and when it was mixed with one of test <strong>in</strong>gredients; thus the digestibility of m<strong>in</strong>erals can be negative due<br />

to the presence of <strong>in</strong>terfer<strong>in</strong>g compounds <strong>in</strong> each feed <strong>in</strong>gredient.<br />

Fecal endogenous losses may be estimated us<strong>in</strong>g a l<strong>in</strong>ear regression of the excretion (or ret ention) on the<br />

<strong>in</strong>take, which is equal to the constant of the regression. Kienzle et al. (1998) estimated dietary P requirement for<br />

adult cats based on a factorial procedure. The authors determ<strong>in</strong>ed endogenous fecal loss, endogenous renal<br />

excretion and true digestibility of P. A basal diet conta<strong>in</strong>ed turkey meat (60%) as the major P source, which was<br />

supplemented with P (source unknown) with varied amount of Ca (Ca/P ratios from 1/1-4/1). This procedure,<br />

however, does not estimate the endogenous fecal P excretion accurately. When endogenous fecal P is to be<br />

determ<strong>in</strong>ed as the constant of a l<strong>in</strong>ear regression l<strong>in</strong>e, the P availability (absorption) of the diet has to be constant<br />

throughout, i.e., constant regression coefficient.<br />

True digestibility by differential approaches<br />

The true digestibility of P may be determ<strong>in</strong>ed without know<strong>in</strong>g the endogenous excretion. Steggerda & Mitchell<br />

(1939) determ<strong>in</strong>ed the true availability of Ca <strong>in</strong> human diets by giv<strong>in</strong>g the test materials at two levels of <strong>in</strong>take, one<br />

very low and the other at about ma<strong>in</strong>tenance, and calculat<strong>in</strong>g the change <strong>in</strong> Ca retention relative to the change <strong>in</strong> Ca<br />

<strong>in</strong>take (i.e., True availability = ∆Retention/∆Intake). A similar approach has been reported by Ammerman et al.<br />

(1957) who determ<strong>in</strong>ed the true absorption percentages of various P compounds by us<strong>in</strong>g the follow<strong>in</strong>g equation;<br />

True absorption (%) = {(Total P <strong>in</strong>take – P <strong>in</strong> basal ration) – (Total P exc. – Basal P exc.)}*100/(Total P <strong>in</strong>take – P <strong>in</strong><br />

basal ration). This formula may be simplified as True absorption = ∆Absorption/∆Intake. In this experiment, all<br />

lambs received a basal ration conta<strong>in</strong><strong>in</strong>g 0.032%P for 4 weeks, and the fecal collections of 5-7d-duration were made<br />

dur<strong>in</strong>g the latter part of the 4wk-depletion period. The supplemental feed<strong>in</strong>g of P lasted for 12d dur<strong>in</strong>g which four<br />

3-day collections were made. The supplemental rations conta<strong>in</strong>ed 0.154-0.158%P, which are well below the<br />

dietary requirement. Kleiber (1975) reported an equation to calculate "partial availability" as follows; Partial<br />

availability = (∆Intake – ∆Feces)/ ∆Intake, which is the same as Ammerman's calculation. Hurwitz (1964)<br />

reported a concept of "net P utilization". This method is also similar to the previously reported <strong>in</strong> that the Net P<br />

utilization =(∆P body/∆P <strong>in</strong>take)*100 The slope of abscissa of the dose-response curve determ<strong>in</strong>ed on various P<br />

<strong>in</strong>takes will be (dP body/dP <strong>in</strong>take)*100. The author estimated total body P content from the tibia P-total body P<br />

relationship that had been determ<strong>in</strong>ed separately. The slope was determ<strong>in</strong>ed <strong>in</strong> the l<strong>in</strong>iar regression below the<br />

m<strong>in</strong>imum requirement. It should be noted that the term "net" normally <strong>in</strong>dicates "apparent", but here the author<br />

used the term as "true". Hurwitz et al. (1978) calculated "fractional absorption" to determ<strong>in</strong>e % absorption of a P<br />

supplement at various Ca levels. The calculation is the same as Ammerman et al. (1957). Bondi (1987) reported<br />

a "comparative bal ance technique" to determ<strong>in</strong>e percentage of utilization of a nutrient. The calculation is as<br />

follows: Utilization (%) =∆Balance*100/∆Intake. Oldham & Emmans (1988) briefly discussed about absolute<br />

response (= output/<strong>in</strong>put), <strong>in</strong>cremental response (=∆output/∆Input), marg<strong>in</strong>al effici ency (= slope of the regression<br />

l<strong>in</strong>e), and the differences between dim<strong>in</strong>ish<strong>in</strong>g returns and the broken stick (l<strong>in</strong>e) responses. The differential<br />

method of estimat<strong>in</strong>g the true digestibility or availability is based on the assumption that the endogenous P excretion<br />

is constant at different dietary P <strong>in</strong>takes, the assumption of which has been questioned by Kleiber et al. (1951) as<br />

mentioned above. Ammerman (1995) also discussed this po<strong>in</strong>t by us<strong>in</strong>g terms "M<strong>in</strong>imum endogenous"<br />

(obligatory) and "Variabl e endogenous". When ∆<strong>in</strong>take is very small (i.e., similar levels of <strong>in</strong>take), the total<br />

endogenous will be similar. Thus, the slope method reported by Kleiber and Hurwitz as discussed above is free<br />

from such a compromise, but construction of a best-fit dose-response curve by measur<strong>in</strong>g responses at various P<br />

levels will be essential (thus more laborious than the differential method). The slope method also gives the<br />

availability of a dietary nutrient at various fractions of dietary levels as reported by Gahl et al. (1995, 1996) for<br />

lys<strong>in</strong>e us<strong>in</strong>g pigs and rats. The authors def<strong>in</strong>ed that the marg<strong>in</strong>al efficiency (d retention/d <strong>in</strong>take) is the effici ency<br />

of retention or ga<strong>in</strong> for a small <strong>in</strong>crement of the am<strong>in</strong>o acid added to the diet. The maximum marg<strong>in</strong>al effici ency<br />

occurred at about 40% of the maximum retention of lys<strong>in</strong>e. They concluded that s<strong>in</strong>ce dim<strong>in</strong>ish<strong>in</strong>g returns affect<br />

the upper 55-65% of the response range, this <strong>in</strong>formation has to be considered <strong>in</strong> formulat<strong>in</strong>g economically feasible<br />

diets that would maximize economic returns rather than maximize animal growth. Rodehutscord et al. (2000)<br />

reported similar results on P us<strong>in</strong>g ra<strong>in</strong>bow trout.<br />

© 2000, 2005. Shozo H. Sugiura. All rights reserved.<br />

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