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Influence of Dietary Fibers on Performance and Fermentation Characteristics of Gut Contents from Growing Chicks
Influence of Dietary Fibers on Performance and Fermentation Characteristics of Gut Contents from Growing Chicks S. C. RICKE, P. J. VAN DER AAR, G. C. FAHEY, JR., and L. L. BERGER Department of Animal Science, University of Illinois, Urbana, Illinois 61801 (Received for publication September 18, 1981)
1982 Poultry Science61:1335-1343 INTRODUCTION A l t h o u g h fiber (plant cell wall constituents) present in feedstuffs is resistant t o degradation by digestive enzymes secreted by certain organs of t h e chick, it can be degraded to oligosaccharides and monosaccharides b y e n z y m e s p r o duced b y microorganisms which inhabit t h e gastrointestinal tract. T h e sugar m o n o m e r s are t h e n fermented by bacteria to volatile fatty acids ( V F A ) which serve as energy sources (Hungate, 1 9 6 6 ; Kass et al., 1980). In p o u l t r y , fiber f e r m e n t a t i o n apparently occurs in t h e ceca ( T h o r n b u r n and Willcox, 1 9 6 5 ) , crop (Bayer et al., 1 9 7 8 ) , and lower gastrointestinal tract (Annison et al., 1968). T h e physicochemical properties of fibrous polysaccharides are i m p o r t a n t as t h e chemical c o m p o s i t i o n determines t h e bacterial species t h a t ferments t h e polysaccharides (Bryant, 1 9 7 4 ) , while t h e physical properties d e t e r m i n e
the r e t e n t i o n t i m e of digesta in t h e g u t (Heller et al, 1 9 8 0 ) . T h e longer the r e t e n t i o n t i m e in the ceca and intestine, t h e greater t h e opport u n i t y for microbial degradation of fiber and the greater t h e p r o d u c t i o n of V F A (Hungate, 1966). T h e objectives of these studies were: 1) t o examine t h e effects of fiber source and particle size on performance of chicks, 2 ) , t o e x a m i n e the effects of dietary fibers varying widely in chemical c o m p o s i t i o n on p e r f o r m a n c e of chicks, and 3) t o evaluate the effects of incorporating certain cell wall c o m p o n e n t s i n t o practical corn-soybean meal diets fed t o b o t h 1 and 8-day-old H u b b a r d broilers on p e r f o r m a n c e and V F A c o n c e n t r a t i o n s of gut c o n t e n t s . EXPERIMENTAL PROCEDURE In E x p e r i m e n t s 1 and 2, 8-day-old N e w Hampshire X C o l u m b i a n female chicks were
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ABSTRACT Four experiments were conducted to evaluate the inclusion of fiber from various sources and of various particle sizes in both semipurified and corn-soybean meal diets fed to chicks. Growth rate and feed efficiency were the performance measurements tested. In Experiment 1, 8-day-old New Hampshire X Columbian female chicks were fed polyethylene (PE) and alfalfa cell walls (ACW) of varying particle sizes (<300 Mm, 300 to 600 Mm, and 600 to 1200 Mm), incorporated into diets at the 8% level. A 0% fiber and an 8% solka floe treatment were used as controls. Consumption of ACW resulted in higher (P<.05) growth rates (6.5 vs. 5.4 g/day) and better (P<.05) feed efficiencies (.46 vs. .41) compared with chicks fed PE, regardless of particle size. In Experiment 2, two levels (4 and 8%) of pectin, lignin and xylose:gum arabic (50:50 mix), were tested using 8-day-old chicks of the same strain. Both levels of lignin and xylose:gum arabic (50:50 mix) resulted in superior grains compared to chicks fed control diets, while 8% pectin resulted in lower (P<.05) growth rates (3.3 g/day) than controls (5.2 g/day). Feed efficiencies followed the same pattern as growth rate. In Experiments 3 and 4, 1-day-old chicks (Experiment 3) and 8-dayold chicks (Experiment 4) of the Hubbard strain were fed two levels (5 and 10%) of gum arabic, xylose, and lignin substituted for corn in practical-type diets. Controls included 0% fiber and 5% arenaceous flour treatments. In Experiment 3, chicks fed both levels of xylose and lignin gained better (P<.05) than controls, whereas those fed 5% gum arabic performed poorer (12.3 g/day for controls; 9.6 g/day for gum arabic). Feed efficiency results showed similar trends. Growth rates of 8-day-old chicks fed 10% gum arabic and 10% xylose were lower (P<.05) than controls (28.4 and 30.7 vs. 31.6 g/day, respectively). Only chicks fed 10% gum arabic had lower feed efficiencies than controls. Cecal and large intestinal contents from chicks fed 10% xylose contained the highest concentration of volatile fatty acids (VFA) (83.5 and 79.9 Mmoles/ml). Chicks fed lignin had the least cecal VFA (60.8 Mmoles/ml) while chicks fed lignin and those fed the control diet had the least large intestinal VFA. Acetate was the predominant VFA while propionate was present in the lowest molar concentration. (Key words: chicks, fiber, particle size, performance, volatile fatty acids)
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Chicks were weighed weekly, t h e a m o u n t of feed remaining was recorded, and leftover feed was removed. Water was supplied ad libitum. Parameters measured included growth rate and feed c o n s u m p t i o n . Gain:feed ratios were calculated. In E x p e r i m e n t s 3 and 4, 1-day-old (Experim e n t 3) and 8-day-old male chicks ( E x p e r i m e n t 4 ) of t h e H u b b a r d strain were fed t w o levels (5 and 10%) of gum arabic, xylose, a n d lignin substituted for corn in practical corn-soybean
TABLE 1. Composition of the basal diet fed to chicks in Experiments 1 and 2 Diet constituent Cornstarch Casein Methionine Corn oil a Mineral mix" Vitamin mix c Ethoxyquin
(%) Variable (to 100%) 25.3 .35 10.0 5.4 1.0 125 mg/kg
Mazola corn oil. Composed of each of the following (percent of diet): CaC0 3 , .3; Ca 3 (P0 4 ) 2 , 2 . 8 ; K 2 H P 0 4 , .9; NaCl, .9; MgS0 4 -7H 2 0, .4; MnSO„ -HjO, .07; Fe citrate, .05; ZnC0 3 , .01; C u S 0 4 - 5 H 2 0 , .002; H 3 B 0 3 , .009; N a 2 M o 0 4 ' 2 H 2 0 , .0009; KI, .004; C o S 0 4 - 7 H 2 0 , .0001;Na 2 SeO 3 , .00002. c Composed of each of the following (percent of total vitamin mix): vitamin A palmitate (250,000 IU/g-500,000 IU), .20; vitamin D 3 (400,000 IU/g284,000 IU), .071; DL-a-tocopherol acid succinate (1,210 IU/g-12,000 IU), .99; menadione, .02; riboflavin, .066; calcium pantothenate, .25; niacin, .4; vitamin B, 2 ticturate, .00044; folic acid, .02; biotin, .002; ascorbic acid, 3.00; pyridoxine-HCl, .025; thiamine-HC1, .025; choline CI, 22.00; powdered starch, 72.93.
meal diets (Table 2 ) . C o n t r o l s included 0% fiber and 5% arenaceous flour t r e a t m e n t s . In
TABLE 2. Composition of the basal diet fed to chicks in Experiments 3 and 4
Diet constituent Corn (8.13% crude protein) Soybean meal (48.31% crude protein) Corn gluten meal (60.00% crude protein) Alfalfa meal (17.00% crude protein) Corn oil a Dicalcium phosphate Limestone Salt MnSO„ (28% Mn) Choline chloride Vitamin mix D ZnC0 3 Antibiotic premix (chlortetracycline)
International feed number 4-02-935 5-04-594 5-02-900 1-00-023 4-97-882 6-01-080 6-02-632
(%) 52.17 38.83 2.00 1.00 2.00 2.20 1.00 .05 .05 .10 .10 .01 .05
Mazola corn oil. Composed of each of the following (percent of total vitamin mix): Micro A/D (500,000 IU/250,000 IU), .40; vitamin E (200 IU/g — Rovimix), 4.54; Hetrazeen (vitamin K), .25; riboflavin, .36; niacin, 2.70; calcium pantothenate, 1.10; vitamin B 12 ticturate (1 mg/g), .90;biotin, .01;soybean meal (fine grind), 89.74.
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allotted in a completely r a n d o m i z e d design with 3 p e n s of 5 chicks per t r e a t m e n t . Petersime b r o o d e r s ( 3 3 C) placed in an environmentally controlled r o o m (23 C) were used t o h o u s e t h e chicks. T h e basal diet (Table 1) was t h e same for b o t h e x p e r i m e n t s . T r e a t m e n t s in E x p e r i m e n t 1 included three different particle sizes of p o l y e t h y l e n e (PE) and alfalfa cell walls (ACW) ( < 3 0 0 ^ m , 3 0 0 t o 6 0 0 jum, 6 0 0 t o 1 2 0 0 jiim) included in t h e diet at the 8% level with a 0% fiber control and an 8% solka floe positive control. Particles were separated using a dry-sieving t e c h n i q u e (Ro-tap p o r t a b l e sieve shaker m o d e l R X - 2 4 ) . Alfalfa cell walls were prepared as described b y Goering and Van Soest ( 1 9 7 0 ) . T r e a t m e n t s in E x p e r i m e n t 2 included t w o levels (4 and 8%) of pectin, lignin and xylose:gum arabic ( 5 0 : 5 0 m i x ) , with 0% fiber and 8% solka floe t r e a t m e n t s used as controls.
FIBER EFFECTS ON CHICKS
RESULTS Experiment 1. Performance data for chicks in Experiment 1 are summarized in Table 3. Chicks fed 8% solka floe, PE (300 to 600 Mm), and all ACW treatments gained more (P<.05) than those fed the 0% fiber basal. However, this improvement was not noted in feed conversion, although chicks fed ACW tended to have higher gainrfeed ratios than those chicks fed either of the control diets (basal or 8% solka floe). Chicks fed the ACW treatments gained more, ingested more feed, and converted feed to gain more efficiently than chicks fed PE-containing diets, regardless of particle size. The 300 to 600 /Urn particle size appeared to be optimal as improvements in gain were noted for both PE and ACW-fed chicks when compared to chicks fed either of the other two particle sizes (<300 jum and 600 to 1200 Mm). The highest feed efficiencies were also observed for chicks fed PE and ACW diets of this same particle size. Experiment 2. Table 4 shows performance data for chicks fed different fiber sources. Type of fibrous carbohydrate influenced chick performance as compared to chicks fed diets not containing fiber. Chicks fed 8% solka floe showed no improvement in gain or feed efficiency over basal-fed chicks. Chicks fed 8% pectin gained poorly and were the poorest con-
TABLE 3. Performance of chicks in Experiment 1 (particle size)3-
Gain
Treatment 1. 2. 3. 4. 5. 6. 7. 8.
Basal Solka flocd Polyethylene (<300 Mm) Polyethylene (300 to 600 jum) Polyethylene (600 to 1200 Mm) Alfalfa cell walls (<300 Mm) Alfalfa cell walls (300 to 600 Mm) Alfalfa cell walls (600 to 1200 Mm)
SEMe
(g/day)t>c 5.1 5.5 5,1 5.8 5.2 6.5 6.9 6.3
Feed intake (g/day)c 12.1 13.3 12.2 13.6 13.0 14.6 14.4 13.8
Gain/ feed c .42 .42 .41 .43 .40 .44 .48 .45 .03
Means of triplicate groups of 5 chicks during the period 8 to 22 days posthatching. Basal different (P<.05) from average of all other treatments. c Polyethylene treatments differ (P<.05) from alfalfa cell wall treatments. Brown Co., Des Plaines, IL. Chemical composition (percentage): cellulose, 78.9; hemicellulose, 10.4; acid detergent lignin, 3.0. Standard error of the mean.
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these trials, we wished to determine if fiber consumption affected the nutritional status of newly hatched chicks. Also, we wished to determine which carbohydrate fraction of the xylose:gum arabic mixture, if any, resulted in the improved performance of chicks noted in earlier trials; therefore, xylose and gum arabic were each fed separately. Allotment, feeding, weighing schedules, and performance measurements tested were essentially the same as in Experiments 1 and 2 except that 3 chicks per pen were used in Experiment 3. At the termination of Experiment 4, chicks were taken off feed for 2 hr after which time they were sacrificed and total cecal and large intestinal contents collected for VFA determinations. Total contents were centrifuged at 0 C immediately after collection and the supernatant prepared for gas liquid chromatographic analyses according to Erwin et al. (1961). The VFA concentrations (g/liter) of acetate, propionate, and butyrate were determined on a Gow Mac Gas Chromatograph 750 equipped with a flame ionization detector. Total VFA concentration (^moles/ml) and molar percentages of each of the three acids were calculated. In all experiments, multiple comparisons between treatments were statistically tested by an F-test at an a = .05 level (Carmer and Swanson, 1973).
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TABLE 4. Performance of chicks in Experiment 2 (fiber source)^
Treatment Control 8%solkaflocf 4% peering 8% peering 4% lignin" 8% lignin" 4% xylose:gum arabic (50:50 mix)' 8% xylose:gum arabic (50:50 mix) 1
5.1b 5.3bc 5.oa b 3.3 a 6.5bc 7.0C
6.5bc 6.5bc
Feed intake (g/day) 14.2 a 14.7 a 18.8 C 21.5 d 15.5ab 16.9 b c 15.2ab 16.3ab
SEMJ
Gain/ feed .36 c .36 c
ai° .15^ .42C .4ic .42 c .39 c .02
' ' ' Means in the same column bearing different superscripts differ (P<.05). f
Means for triplicate groups of 5 chicks during the period 8 to 22 days posthatching. Described in Table 3.
%igma Chemical Co., St. Louis, MO. Partially methoxylated polygalacturonic acid from citrus fruits (without sucrose or other sugars). Westvaco Corp., Charleston, SC. Softwood Kraft lignin, Indulin ATR-CK1, RLX 4290-47. Sigma Chemical Co., St. Louis, MO. A mixture of 50% D(+) xylose and 50% gum arabic, a branched polymer of galactose, rhamnose, arabinose, and glucuronic acid as the calcium, magnesium, and potassium salts with a molecular weight of approximately 250,000. ^Pooled standard error of the mean.
verters of feed t o gain. Chicks fed 8% lignin gained significantly b e t t e r t h a n control-fed chicks. Level of fibrous c a r b o h y d r a t e in t h e diet appeared t o have s o m e effect on feed efficiency b u t n o t o n gain (with t h e exception of those chicks fed 8% p e c t i n ) . Experiment 3. Table 5 summarizes performance data for chicks fed practical cornsoybean meal diets starting on day 1 posthatching. Chicks fed t h e inert bulking agent, arenaceous flour, showed n o response in either gain or feed conversion c o m p a r e d with t h e control. Chicks fed 5% gum arabic had p o o r e r growth rates t h a n controls while those fed 10% gum arabic had lower b u t nonsignificantly different growth rates than controls. Gain: feed ratios were lower ( P < . 0 5 ) for chicks fed b o t h levels of g u m arabic. Chicks fed either level of xylose or lignin grew faster ( P < . 0 5 ) t h a n t h e controls and showed improved ( P < . 0 5 ) feed conversions. Increasing t h e level of fiber in the diet from 5 t o 10% improved ( P < . 0 5 ) gain in xylose-fed chicks b u t had n o effect ( P > . 0 5 ) on lignin-fed chicks. Gain:feed ratios were statistically similar for chicks fed b o t h levels of xylose and lignin since feed in-
t a k e , in b o t h cases, was greater ( P < . 0 5 ) at t h e higher fiber level. Experiment 4. Table 6 summarizes performance data collected in E x p e r i m e n t 4 from chcks fed diets identical t o those of E x p e r i m e n t 3; however, chicks were started at day 8 posthatching. T h e r e was n o i m p r o v e m e n t ( P < . 0 5 ) in gain or feed conversion with cell wall carboh y d r a t e additions. Chicks fed 10% gum arabic or 10% xylose had p o o r e r ( P < . 0 5 ) growth rates t h a n controls. This depression was particularly evident with gum arabic-fed chicks where an increase from 5 t o 10% caused a decrease in gain of 11.8%. A level effect was observed and chicks fed 5% fiber converted feed to gain m o r e efficiently t h a n those fed 10% levels of fiber. Volatile fatty acid concentrations of contents t a k e n from ceca and larger intestine of chicks in E x p e r i m e n t 4 are outlined in Table 7. Total c o n c e n t r a t i o n s (jMmoles/ml) of cecal V F A was highest for chicks fed 10% xylose. However, n o statistical differences in cecal V F A c o n c e n t r a t i o n s were n o t e d between basal, 5% arenaceous flour, 10% gum arabic, 5% xylose, or 10% xylose-fed chicks. T h e lowest c o n c e n t r a t i o n of cecal V F A occurred w h e n
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1. 2. 3. 4. 5. 6. 7. 8.
Gain (g/day)
FIBER EFFECTS ON CHICKS
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TABLE 5. Performance of chicks in Experiment 3 (l-day-old chicks)^ Feed intake (g/day)
Gain/ feed
Basal 5% arenaceous flour' 5% gum arabicg 10% gum arabicg 5% xyloseh 10% xylose" 5% lignin1 8. 10%lignini
12.0 a 12.6 a 9.6 b 11.la 14.2<= 17.2 d 16.9 d 16.6 d
16.7 a 17.8 a 15.9 a 18.9 a 18.5 a 21.3b 19.7 a 21.7 b
.72 a .71a .61b .59 b .77 c .81c .86 c .77=
SEMJ
± .5
± .7
a
f
+ .02
' ' ' Means in the same column bearing different superscripts differ (P<.05). Means of triplicate groups of 3 chicks during the period 1 to 14 days posthatching.
Wedron Silica Division, Park Ridge, IL. Chemical composition (percentage): SiO, 99.88; F e , 0 3 , .02; A1 2 0 3 , .10;TiO 3 , .015; CaO, .015;andMgO, .005. ^Described in Table 4. Described in Table 4. Described in Table 4. ^Pooled standard error of the mean.
TABLE 6. Performance of chicks in Experiment 4 (8-day-old cbicks)e
Treatment
Gain (g/day)
Feed intake (g/day)
Gain/ feed
1. 2. 3. 4. 5. 6. 7. 8.
32.3 C 30.9 b c 32.2 C 28.4 a 30.7abc 28.6 a b 31.6 C 30.6abc
47 gabc 50.9 b c 49 4abc 53.5 C 45.6 a 45.8 a 50.2abc 51.9 b c
.67d .61bc .64 c d .53a .67d .63bcd .63 b c d .59b
± 1.6
.02
Basal 5% arenaceous flour? 5% gum arabicg 10% gum arabicg 5% xylose" 10% xylose" 5% lignin' 10% lignin1
SEMJ
a,b,c,d Means in the same column bearing different superscripts differ (P<.05). f
Means of triplicate groups of 5 chicks during the period 8 to 22 days posthatching. Described in Table 5.
^Described in Table 4. Described in Table 4. 'Described in Table 4. ••Pooled standard error of the mean.
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Treatment
Gain (g/day)
Basal 5% arenaceous flour^ 5% gum arabicS 10% gum arabicg 5% xylose" 10% xylose" 5% lignini 10% lignini
± 1.4
70.5a 72.9ab 81.0de 84.4 e 77_ 4 cd 78.4cd 76.3bc 76.3bc
82.lc 78.9 b c 65.4ab 70.4abc 71.3abc 83.5 C 60.7a 60.8 a
5.0
(Ac)
(/umoles/ml)
CecuiTl
'Described in Table 5.
Pooled standard error of the mean.
•Not detectable.
J
'Described in Table 4.
Described in Table 4.
"Described in Table 4.
1.3
20.7C 19.0 C 13.2 b 8.9 a 14.5 b 14.1b 14.7 b 13.4 b
8.8bc g jabc 5.9 a 6.6ab 8.0abc 7.4 a b 7.2ab 10.4 C .8
(Bu)
(Pr)
Molar %
a.b.^d.e,Means in the same column bearing different superscripts differ (P<.05).
SEMJ
1. 2. 3. 4. 5. 6. 7. 8.
Treatment
VFA con-
5.9
21. l a 23.3 a 43.5b 63.5cd 58.3bc 79.9 d 24.6 a 25.6 a
VFA concentration (Mmoles/ml)
TABLE 7. Total concentrations (iimoles/ml) and molar percentages of VFA s in cecal and l of chicks in Experiment 4
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FIBER EFFECTS ON CHICKS
DISCUSSION
Although fiber particle size did not influence (P<.05) performance of chicks fed semipurified diets, those fed the intermediate size particles (300 to 600 /xm) tended to have the highest rates of gain and feed efficiencies. This effect was noted with both bulking agents (PE and ACW), suggesting that particle size may be an important property of a fibrous ingredient, regardless of whether the fiber is potentially fermentable (ACW) or not (PE). Scott and Forbes (1958) noted that dietary fiber improved performance of chicks by simply narrowing the protein: calorie ratio, thus causing the chick to eat more of the most limiting amino acids to meet its energy needs. Results obtained with solka floe and PE-fed (300 to 600 urn and 600 to 1200 Urn) chicks in our studies support this work. Chicks fed ACW, however, performed better than those fed either solka floe or PE, indicating that possibly some component of ACW or microbial fermentation of the structural carbohydrates present in ACW may have resulted in beneficial effects on performance of the chick. This hypothesis is supported by data collected
in Experiment 2 (Table 4) where it was noted that certain of the components of ACW resulted in improvements in growth rates of chicks. However, chicks fed pectin, especially at the 8% level, performed poorly, as evidenced by low growth rates and feed efficiencies. Pectin-fed chicks ingested more (P<.05) feed than those fed other fiber types and excreted feces with a sticky consistency. The latter observation is supported by work of Wagner and Thomas (1977). These workers suggested that the growth depression seen with pectin was due to the growth of adverse intestinal microorganisms. In a subsequent paper, Wagner and Thomas (1978) suggested that a penicillinsensitive, spore-forming organism, which produced large amounts of gas and butyric acid fermentatively, depressed growth of chicks fed pectin. In certain instances, the feeding of D-xylose and L-arabinose, two primary components of hemicellulose, resulted in depressed growth rates of chicks (Wagh and Waibel, 1966; Baker, 1977), which may be attributed to direct competition with glucose for absorption sites (Ross et al, 1972) or poor metabolizability of these sugars (Wagh and Waibel, 1967). Wagh and Waibel (1966) found no significant differences between treatments when 10% xylosefed chicks were compared with controls. However, Baker (1977) found significant depressions at higher levels of xylose feeding. Very little work has been conducted on the nutritional value of the purified Kraft wood lignin used in this work. Most research to date has centered on its adsorptive properties (Kay et al, 1979; Rubio et al, 1979). Feeding both 4 and 8% lignin to chicks resulted in higher (P<.05) growth rates compared to control-fed chicks. Increased feed intake is a possible causal factor in the response, but it has recently been shown by Zemek et al. (1979) that certain phenolic constituents of lignin have antibioticlike properties. Also, Miller et al. (1979) found that various fractions isolated from wood hemicellulose extract, a chemically complex material high in phenolics, resulted in improvements in performance of chicks when incorporated into practical corn-soybean meal diets. Further evidence that phenolic compounds are involved in the growth-promoting properties of plant extracts can be noted in the work of Johanning and O'Dell (1981). They found that the cell wall material from cabbage and alfalfa which promoted guinea pig performance inhibited
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5 or 10% lignin was fed to chicks. Molar percent acetate was highest in ceca of chicks fed either level of gum arabic and lowest for basal and arenaceous flour-fed chicks. Chicks fed 10% lignin had the highest molar percent propionate (10.4), while those fed 5% gum arabic had the lowest (5.9). Molar percent butyrate was highest in ceca of chicks fed basal and 5% arenaceous flour diets. In the large intestine of chicks, VFA concentrations were low for basal, 5% arenaceous flour, and 5 and 10% lignin-fed chicks. The highest concentrations of VFA in the large intestine occurred when chicks were fed diets containing 10% gum arabic and 10% xylose. Level effects were noted for gum arabic and xylose. Molar percentages of acetate were high while molar percentages of propionate and butyrate were low. Ten percent gum arabic and 5 and 10% xylose-fed chicks had nondetectable levels of propionate. The highest molar percent propionate was noted for 10% lignin-fed chicks. Molar percent butyrate was highest in large intestinal contents of 5 and 10% gum arabicfed chicks and lowest in intestinal contents of chicks fed 10% xylose. No fiber level effect on molar percent was noted.
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Chicks started on cell wall carbohydratesupplemented diets at day 8 posthatching performed differently than those started at day 1. Chicks fed all treatment diets performed equivalent to or poorer than controls. A possible explanation for the different responses due to age may be that Hubbard chicks started at 8 days of age ingest such great quantities of feed (40 to 50 g/day) that rate of passage is increased to the extent that gut microbes do not have time to effectively ferment cell wall carbohydrate arriving at the ceca and large intestine. Adverse intestinal organisms may then compete with the normal flora, resulting in poorer animal performance. This is analogous to the situation which occurs with pectin-fed chicks (Wagner and Thomas, 1977).
Volatile fatty acid data suggested that different cell wall constituents affected gut microbial metabolism. Cecal VFA concentrations were highest for chicks fed basal and 10% xylose diets and lowest for chicks fed lignincontaining diets. Chicks fed 10% lignin had the highest molar percent cecal propionate, indicating that the cecal microbes of chicks fed this fiber source had a different fermentation pattern than those fed the other diets. Since phenolic components of lignin have been shown to have antibiotic-like properties (Zemek et al., 1979), this fermentation pattern may be of significance to the avian species. Much lower VFA concentrations (compared with cecal concentrations) were noted in the large intestine of chicks fed control diets and lignin-containing diets. Chicks fed 10% gum arabic and 10% xylose had the highest concentrations of large intestinal VFA's. Acetate was present in the greatest molar percent while molar percent butyrate was variable among treatments. Propionate was present in the lowest concentration. Chicks fed 10% lignin had the lowest molar percent acetate and the highest molar percent propionate, indicating a similar fermentation pattern to that of the ceca. In conclusion, it appears that cell wall constituents can have striking effects on avian performance and lower gut VFA patterns. More research is needed to define the chemical and physical properties of complex carbohydrates which will result in maximum performance of poultry. REFERENCES Annison, E. T., K. J. Hill, and R. Kenworthy, 1968. Volatile fatty acids in the digestive tract of the fowl. Br. J. Nutr. 22:207-216. Baker, D. H., 1977. Xylose and xylan utilization by the chick. Poultry Sci. 56:2105-2107. Bayer, R. C, W. H. Hoover, and F. V. Muir, 1978. Dietary fiber and meal feeding influence on broiler growth and crop fermentation. Poultry Sci. 57:1456-1459. Bryant, M. P., 1974. Nutritional features and ecology of predominant anaerobic bacteria of the intestinal tract. Am. J. Clin. Nutr. 27:1313-1319. Carmer, S. G., and M. R. Swanson, 1973. An evaluation of ten pairwise multiple comparisons procedures by Monte Carlo methods. J. Am. Stat. Assoc. 68:66-74. Erwin, E. S., G. J. Marco, and E. M. Emery, 1961. Volatile fatty acid analysis of blood and rumen fluid by gas chromatography. J. Dairy Sci. 44: 1768-1771. Goering, H. K., and P. J. Van Soest, 1970. Forage fiber analysis. ARS, USDA Handbook 379. Agric. Res. Serv., Washington, DC.
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growth of an unidentified anaerobic bacteria from guinea pig ceca. Extraction of the cell wall material with alkali resulted in the loss of pcoumaric and caffeic acids, two phenolics common to many plants (Hermann, 1980), and eliminated the antibacterial activity of the cell walls. Both of these acids are components of lignin. The best performances of 1-day-old chicks (Experiment 3) occurred when xylose and lignin were fed, suggesting that chicks fed these carbohydrates evidently establish a favorable gastrointestinal tract environment for utilization of the diet. Since a gut microbial population is established early in life, it appears that feeding certain cell wall constituents such as xylose and ball milled lignin from day 1 of age may result in an altered fermentation by gut microbes as evidenced by changes in the quantity and proportion of volatile fatty acids. Another possible explanation for the improvement in performance due to xylose supplementation may be that the specific activities of the enzymes catalyzing the reactions in the liver where xylose is converted to glucose are increased and thus xylose is being used as an energy source. The former explanation is probably more realistic, however, since even if lignin was degraded to its phenolic constituents and subsequently absorbed, no energy contribution to the animal would be possible. Also, since gum arabic was not well utilized by 1-dayold chicks (as evidenced by poor growth rates and feed efficiencies), it appears that the improvements due to the addition of cell wall carbohydrates is caused by a change in a microbial metabolism in the GI tract rather than a change in postabsorptive (tissue) metabolism.
FIBER EFFECTS ON CHICKS
2419. Rubio, M. A., S. Ingemar Falkehag, B. A. Pethica, and P. Zuman, 1979. The interactions of carcinogens and co-carcinogens with lignin and other components of dietary fiber. Pages 251—271 in Dietary fibers: Chemistry and nutrition. G. E. Inglett and S. Ingemar Falkehag, ed. Academic Press, New York, NY. Scott, H. M., and R. M. Forbes, 1958. The arginine requirement of chicks in relation to diet composition. Poultry Sci. 37:1347-1349. Thornburn, C. C , and J. S. Willcox, 1965. The caeca of the domestic fowl and digestion of the crude fiber complex. II. Experiments in vivo with fistulated birds, and the artificial and isolated caecum in vitro. Br. Poultry Sci. 6:33—43. Wagh, P. V., and P. E. Waibel, 1966. Metabolizability and nutritional implications of L-arabinose and D-xylose for chicks. J. Nutr. 9 0 : 2 0 7 - 2 1 1 . Wagh, P. V., and P. E. Waibel, 1967. Metabolism of L-arabinose and D-xylose by chicks. J. Nutr. 92: 491-496. Wagner, D. D., and O. P. Thomas, 1977. A rye type growth depression of chicks fed pectin. Poultry Sci. 56:615-619. Wagner, D. D., and O. P. Thomas, 1978. Influence of diets containing rye or pectin on the intestinal flora of chicks. Poultry Sci. 57:971-975. Zemek, J., B. Kosikova, J. Augustin, and D. Joniak, 1979. Antibiotic properties of lignin components. Folia Microbiol. 24:483-486.
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Heller, S. N„ L. R. Hackler, J. M. Rivers, P. J. Van Soest, D. A. Roe, B. A. Lewis, and J. B. Robertson, 1980. Dietary fiber: the effect of particle size of wheat bran on colonic function of young adult men. Am..J. Clin. Nutr. 33:1734-1744. Hermann, K., 1980. Pflanzenphenole als Inhaltsstoffe von Lebensmitteln Ernaehr. Umsch. 27: 75-80. Hungate, R. E., 1966. The Rumen and Its Microbes. Academic Press, New York, NY. Johanning, G. L., and B. L. O'Dell, 1981. Inhibition of a cecal anaerobe by a dietary fiber component. Fed. Proc. 40:485. (Abstr.) Kass, M. L., P. J. Van Soest, and W. G. Pond. 1980. Utilization of dietary fiber from alfalfa by growing swine. II. Volatile fatty acid concentrations in and disappearance from the gastrointestinal tract. J. Anim. Sci. 50:192-197. Kay, R. M., S. M. Strasberg, C. N. Petrunka, and M. Wayman, 1979. Differential absorption of bile acids by lignins. Pages 57—65 in Dietary Fibers: Chemistry and Nutrition. G. E. Inglett and S. Ingemar Falkenhag, ed. Academic Press, New York, NY. Miller, B. L., G. C. Fahey, Jr., and S. L. Schussler, 1979. Isolation of biologically active fractions from wood hemicellulose extracts. J. Anim. Sci. 48:1129-1134. Ross, A., A. W. Rubin, and J. J. Deren, 1972. Differential permeability of the proximal and distal rabbit small bowel. J. Clin. Invest. 51:2414—
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1982 Poultry Science61:1335-1343 INTRODUCTION A l t h o u g h fiber (plant cell wall constituents) present in feedstuffs is resistant t o degradation by digestive enzymes secreted by certain organs of t h e chick, it can be degraded to oligosaccharides and monosaccharides b y e n z y m e s p r o duced b y microorganisms which inhabit t h e gastrointestinal tract. T h e sugar m o n o m e r s are t h e n fermented by bacteria to volatile fatty acids ( V F A ) which serve as energy sources (Hungate, 1 9 6 6 ; Kass et al., 1980). In p o u l t r y , fiber f e r m e n t a t i o n apparently occurs in t h e ceca ( T h o r n b u r n and Willcox, 1 9 6 5 ) , crop (Bayer et al., 1 9 7 8 ) , and lower gastrointestinal tract (Annison et al., 1968). T h e physicochemical properties of fibrous polysaccharides are i m p o r t a n t as t h e chemical c o m p o s i t i o n determines t h e bacterial species t h a t ferments t h e polysaccharides (Bryant, 1 9 7 4 ) , while t h e physical properties d e t e r m i n e
the r e t e n t i o n t i m e of digesta in t h e g u t (Heller et al, 1 9 8 0 ) . T h e longer the r e t e n t i o n t i m e in the ceca and intestine, t h e greater t h e opport u n i t y for microbial degradation of fiber and the greater t h e p r o d u c t i o n of V F A (Hungate, 1966). T h e objectives of these studies were: 1) t o examine t h e effects of fiber source and particle size on performance of chicks, 2 ) , t o e x a m i n e the effects of dietary fibers varying widely in chemical c o m p o s i t i o n on p e r f o r m a n c e of chicks, and 3) t o evaluate the effects of incorporating certain cell wall c o m p o n e n t s i n t o practical corn-soybean meal diets fed t o b o t h 1 and 8-day-old H u b b a r d broilers on p e r f o r m a n c e and V F A c o n c e n t r a t i o n s of gut c o n t e n t s . EXPERIMENTAL PROCEDURE In E x p e r i m e n t s 1 and 2, 8-day-old N e w Hampshire X C o l u m b i a n female chicks were
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ABSTRACT Four experiments were conducted to evaluate the inclusion of fiber from various sources and of various particle sizes in both semipurified and corn-soybean meal diets fed to chicks. Growth rate and feed efficiency were the performance measurements tested. In Experiment 1, 8-day-old New Hampshire X Columbian female chicks were fed polyethylene (PE) and alfalfa cell walls (ACW) of varying particle sizes (<300 Mm, 300 to 600 Mm, and 600 to 1200 Mm), incorporated into diets at the 8% level. A 0% fiber and an 8% solka floe treatment were used as controls. Consumption of ACW resulted in higher (P<.05) growth rates (6.5 vs. 5.4 g/day) and better (P<.05) feed efficiencies (.46 vs. .41) compared with chicks fed PE, regardless of particle size. In Experiment 2, two levels (4 and 8%) of pectin, lignin and xylose:gum arabic (50:50 mix), were tested using 8-day-old chicks of the same strain. Both levels of lignin and xylose:gum arabic (50:50 mix) resulted in superior grains compared to chicks fed control diets, while 8% pectin resulted in lower (P<.05) growth rates (3.3 g/day) than controls (5.2 g/day). Feed efficiencies followed the same pattern as growth rate. In Experiments 3 and 4, 1-day-old chicks (Experiment 3) and 8-dayold chicks (Experiment 4) of the Hubbard strain were fed two levels (5 and 10%) of gum arabic, xylose, and lignin substituted for corn in practical-type diets. Controls included 0% fiber and 5% arenaceous flour treatments. In Experiment 3, chicks fed both levels of xylose and lignin gained better (P<.05) than controls, whereas those fed 5% gum arabic performed poorer (12.3 g/day for controls; 9.6 g/day for gum arabic). Feed efficiency results showed similar trends. Growth rates of 8-day-old chicks fed 10% gum arabic and 10% xylose were lower (P<.05) than controls (28.4 and 30.7 vs. 31.6 g/day, respectively). Only chicks fed 10% gum arabic had lower feed efficiencies than controls. Cecal and large intestinal contents from chicks fed 10% xylose contained the highest concentration of volatile fatty acids (VFA) (83.5 and 79.9 Mmoles/ml). Chicks fed lignin had the least cecal VFA (60.8 Mmoles/ml) while chicks fed lignin and those fed the control diet had the least large intestinal VFA. Acetate was the predominant VFA while propionate was present in the lowest molar concentration. (Key words: chicks, fiber, particle size, performance, volatile fatty acids)
RICKE ET AL.
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Chicks were weighed weekly, t h e a m o u n t of feed remaining was recorded, and leftover feed was removed. Water was supplied ad libitum. Parameters measured included growth rate and feed c o n s u m p t i o n . Gain:feed ratios were calculated. In E x p e r i m e n t s 3 and 4, 1-day-old (Experim e n t 3) and 8-day-old male chicks ( E x p e r i m e n t 4 ) of t h e H u b b a r d strain were fed t w o levels (5 and 10%) of gum arabic, xylose, a n d lignin substituted for corn in practical corn-soybean
TABLE 1. Composition of the basal diet fed to chicks in Experiments 1 and 2 Diet constituent Cornstarch Casein Methionine Corn oil a Mineral mix" Vitamin mix c Ethoxyquin
(%) Variable (to 100%) 25.3 .35 10.0 5.4 1.0 125 mg/kg
Mazola corn oil. Composed of each of the following (percent of diet): CaC0 3 , .3; Ca 3 (P0 4 ) 2 , 2 . 8 ; K 2 H P 0 4 , .9; NaCl, .9; MgS0 4 -7H 2 0, .4; MnSO„ -HjO, .07; Fe citrate, .05; ZnC0 3 , .01; C u S 0 4 - 5 H 2 0 , .002; H 3 B 0 3 , .009; N a 2 M o 0 4 ' 2 H 2 0 , .0009; KI, .004; C o S 0 4 - 7 H 2 0 , .0001;Na 2 SeO 3 , .00002. c Composed of each of the following (percent of total vitamin mix): vitamin A palmitate (250,000 IU/g-500,000 IU), .20; vitamin D 3 (400,000 IU/g284,000 IU), .071; DL-a-tocopherol acid succinate (1,210 IU/g-12,000 IU), .99; menadione, .02; riboflavin, .066; calcium pantothenate, .25; niacin, .4; vitamin B, 2 ticturate, .00044; folic acid, .02; biotin, .002; ascorbic acid, 3.00; pyridoxine-HCl, .025; thiamine-HC1, .025; choline CI, 22.00; powdered starch, 72.93.
meal diets (Table 2 ) . C o n t r o l s included 0% fiber and 5% arenaceous flour t r e a t m e n t s . In
TABLE 2. Composition of the basal diet fed to chicks in Experiments 3 and 4
Diet constituent Corn (8.13% crude protein) Soybean meal (48.31% crude protein) Corn gluten meal (60.00% crude protein) Alfalfa meal (17.00% crude protein) Corn oil a Dicalcium phosphate Limestone Salt MnSO„ (28% Mn) Choline chloride Vitamin mix D ZnC0 3 Antibiotic premix (chlortetracycline)
International feed number 4-02-935 5-04-594 5-02-900 1-00-023 4-97-882 6-01-080 6-02-632
(%) 52.17 38.83 2.00 1.00 2.00 2.20 1.00 .05 .05 .10 .10 .01 .05
Mazola corn oil. Composed of each of the following (percent of total vitamin mix): Micro A/D (500,000 IU/250,000 IU), .40; vitamin E (200 IU/g — Rovimix), 4.54; Hetrazeen (vitamin K), .25; riboflavin, .36; niacin, 2.70; calcium pantothenate, 1.10; vitamin B 12 ticturate (1 mg/g), .90;biotin, .01;soybean meal (fine grind), 89.74.
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allotted in a completely r a n d o m i z e d design with 3 p e n s of 5 chicks per t r e a t m e n t . Petersime b r o o d e r s ( 3 3 C) placed in an environmentally controlled r o o m (23 C) were used t o h o u s e t h e chicks. T h e basal diet (Table 1) was t h e same for b o t h e x p e r i m e n t s . T r e a t m e n t s in E x p e r i m e n t 1 included three different particle sizes of p o l y e t h y l e n e (PE) and alfalfa cell walls (ACW) ( < 3 0 0 ^ m , 3 0 0 t o 6 0 0 jum, 6 0 0 t o 1 2 0 0 jiim) included in t h e diet at the 8% level with a 0% fiber control and an 8% solka floe positive control. Particles were separated using a dry-sieving t e c h n i q u e (Ro-tap p o r t a b l e sieve shaker m o d e l R X - 2 4 ) . Alfalfa cell walls were prepared as described b y Goering and Van Soest ( 1 9 7 0 ) . T r e a t m e n t s in E x p e r i m e n t 2 included t w o levels (4 and 8%) of pectin, lignin and xylose:gum arabic ( 5 0 : 5 0 m i x ) , with 0% fiber and 8% solka floe t r e a t m e n t s used as controls.
FIBER EFFECTS ON CHICKS
RESULTS Experiment 1. Performance data for chicks in Experiment 1 are summarized in Table 3. Chicks fed 8% solka floe, PE (300 to 600 Mm), and all ACW treatments gained more (P<.05) than those fed the 0% fiber basal. However, this improvement was not noted in feed conversion, although chicks fed ACW tended to have higher gainrfeed ratios than those chicks fed either of the control diets (basal or 8% solka floe). Chicks fed the ACW treatments gained more, ingested more feed, and converted feed to gain more efficiently than chicks fed PE-containing diets, regardless of particle size. The 300 to 600 /Urn particle size appeared to be optimal as improvements in gain were noted for both PE and ACW-fed chicks when compared to chicks fed either of the other two particle sizes (<300 jum and 600 to 1200 Mm). The highest feed efficiencies were also observed for chicks fed PE and ACW diets of this same particle size. Experiment 2. Table 4 shows performance data for chicks fed different fiber sources. Type of fibrous carbohydrate influenced chick performance as compared to chicks fed diets not containing fiber. Chicks fed 8% solka floe showed no improvement in gain or feed efficiency over basal-fed chicks. Chicks fed 8% pectin gained poorly and were the poorest con-
TABLE 3. Performance of chicks in Experiment 1 (particle size)3-
Gain
Treatment 1. 2. 3. 4. 5. 6. 7. 8.
Basal Solka flocd Polyethylene (<300 Mm) Polyethylene (300 to 600 jum) Polyethylene (600 to 1200 Mm) Alfalfa cell walls (<300 Mm) Alfalfa cell walls (300 to 600 Mm) Alfalfa cell walls (600 to 1200 Mm)
SEMe
(g/day)t>c 5.1 5.5 5,1 5.8 5.2 6.5 6.9 6.3
Feed intake (g/day)c 12.1 13.3 12.2 13.6 13.0 14.6 14.4 13.8
Gain/ feed c .42 .42 .41 .43 .40 .44 .48 .45 .03
Means of triplicate groups of 5 chicks during the period 8 to 22 days posthatching. Basal different (P<.05) from average of all other treatments. c Polyethylene treatments differ (P<.05) from alfalfa cell wall treatments. Brown Co., Des Plaines, IL. Chemical composition (percentage): cellulose, 78.9; hemicellulose, 10.4; acid detergent lignin, 3.0. Standard error of the mean.
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these trials, we wished to determine if fiber consumption affected the nutritional status of newly hatched chicks. Also, we wished to determine which carbohydrate fraction of the xylose:gum arabic mixture, if any, resulted in the improved performance of chicks noted in earlier trials; therefore, xylose and gum arabic were each fed separately. Allotment, feeding, weighing schedules, and performance measurements tested were essentially the same as in Experiments 1 and 2 except that 3 chicks per pen were used in Experiment 3. At the termination of Experiment 4, chicks were taken off feed for 2 hr after which time they were sacrificed and total cecal and large intestinal contents collected for VFA determinations. Total contents were centrifuged at 0 C immediately after collection and the supernatant prepared for gas liquid chromatographic analyses according to Erwin et al. (1961). The VFA concentrations (g/liter) of acetate, propionate, and butyrate were determined on a Gow Mac Gas Chromatograph 750 equipped with a flame ionization detector. Total VFA concentration (^moles/ml) and molar percentages of each of the three acids were calculated. In all experiments, multiple comparisons between treatments were statistically tested by an F-test at an a = .05 level (Carmer and Swanson, 1973).
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TABLE 4. Performance of chicks in Experiment 2 (fiber source)^
Treatment Control 8%solkaflocf 4% peering 8% peering 4% lignin" 8% lignin" 4% xylose:gum arabic (50:50 mix)' 8% xylose:gum arabic (50:50 mix) 1
5.1b 5.3bc 5.oa b 3.3 a 6.5bc 7.0C
6.5bc 6.5bc
Feed intake (g/day) 14.2 a 14.7 a 18.8 C 21.5 d 15.5ab 16.9 b c 15.2ab 16.3ab
SEMJ
Gain/ feed .36 c .36 c
ai° .15^ .42C .4ic .42 c .39 c .02
' ' ' Means in the same column bearing different superscripts differ (P<.05). f
Means for triplicate groups of 5 chicks during the period 8 to 22 days posthatching. Described in Table 3.
%igma Chemical Co., St. Louis, MO. Partially methoxylated polygalacturonic acid from citrus fruits (without sucrose or other sugars). Westvaco Corp., Charleston, SC. Softwood Kraft lignin, Indulin ATR-CK1, RLX 4290-47. Sigma Chemical Co., St. Louis, MO. A mixture of 50% D(+) xylose and 50% gum arabic, a branched polymer of galactose, rhamnose, arabinose, and glucuronic acid as the calcium, magnesium, and potassium salts with a molecular weight of approximately 250,000. ^Pooled standard error of the mean.
verters of feed t o gain. Chicks fed 8% lignin gained significantly b e t t e r t h a n control-fed chicks. Level of fibrous c a r b o h y d r a t e in t h e diet appeared t o have s o m e effect on feed efficiency b u t n o t o n gain (with t h e exception of those chicks fed 8% p e c t i n ) . Experiment 3. Table 5 summarizes performance data for chicks fed practical cornsoybean meal diets starting on day 1 posthatching. Chicks fed t h e inert bulking agent, arenaceous flour, showed n o response in either gain or feed conversion c o m p a r e d with t h e control. Chicks fed 5% gum arabic had p o o r e r growth rates t h a n controls while those fed 10% gum arabic had lower b u t nonsignificantly different growth rates than controls. Gain: feed ratios were lower ( P < . 0 5 ) for chicks fed b o t h levels of g u m arabic. Chicks fed either level of xylose or lignin grew faster ( P < . 0 5 ) t h a n t h e controls and showed improved ( P < . 0 5 ) feed conversions. Increasing t h e level of fiber in the diet from 5 t o 10% improved ( P < . 0 5 ) gain in xylose-fed chicks b u t had n o effect ( P > . 0 5 ) on lignin-fed chicks. Gain:feed ratios were statistically similar for chicks fed b o t h levels of xylose and lignin since feed in-
t a k e , in b o t h cases, was greater ( P < . 0 5 ) at t h e higher fiber level. Experiment 4. Table 6 summarizes performance data collected in E x p e r i m e n t 4 from chcks fed diets identical t o those of E x p e r i m e n t 3; however, chicks were started at day 8 posthatching. T h e r e was n o i m p r o v e m e n t ( P < . 0 5 ) in gain or feed conversion with cell wall carboh y d r a t e additions. Chicks fed 10% gum arabic or 10% xylose had p o o r e r ( P < . 0 5 ) growth rates t h a n controls. This depression was particularly evident with gum arabic-fed chicks where an increase from 5 t o 10% caused a decrease in gain of 11.8%. A level effect was observed and chicks fed 5% fiber converted feed to gain m o r e efficiently t h a n those fed 10% levels of fiber. Volatile fatty acid concentrations of contents t a k e n from ceca and larger intestine of chicks in E x p e r i m e n t 4 are outlined in Table 7. Total c o n c e n t r a t i o n s (jMmoles/ml) of cecal V F A was highest for chicks fed 10% xylose. However, n o statistical differences in cecal V F A c o n c e n t r a t i o n s were n o t e d between basal, 5% arenaceous flour, 10% gum arabic, 5% xylose, or 10% xylose-fed chicks. T h e lowest c o n c e n t r a t i o n of cecal V F A occurred w h e n
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1. 2. 3. 4. 5. 6. 7. 8.
Gain (g/day)
FIBER EFFECTS ON CHICKS
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TABLE 5. Performance of chicks in Experiment 3 (l-day-old chicks)^ Feed intake (g/day)
Gain/ feed
Basal 5% arenaceous flour' 5% gum arabicg 10% gum arabicg 5% xyloseh 10% xylose" 5% lignin1 8. 10%lignini
12.0 a 12.6 a 9.6 b 11.la 14.2<= 17.2 d 16.9 d 16.6 d
16.7 a 17.8 a 15.9 a 18.9 a 18.5 a 21.3b 19.7 a 21.7 b
.72 a .71a .61b .59 b .77 c .81c .86 c .77=
SEMJ
± .5
± .7
a
f
+ .02
' ' ' Means in the same column bearing different superscripts differ (P<.05). Means of triplicate groups of 3 chicks during the period 1 to 14 days posthatching.
Wedron Silica Division, Park Ridge, IL. Chemical composition (percentage): SiO, 99.88; F e , 0 3 , .02; A1 2 0 3 , .10;TiO 3 , .015; CaO, .015;andMgO, .005. ^Described in Table 4. Described in Table 4. Described in Table 4. ^Pooled standard error of the mean.
TABLE 6. Performance of chicks in Experiment 4 (8-day-old cbicks)e
Treatment
Gain (g/day)
Feed intake (g/day)
Gain/ feed
1. 2. 3. 4. 5. 6. 7. 8.
32.3 C 30.9 b c 32.2 C 28.4 a 30.7abc 28.6 a b 31.6 C 30.6abc
47 gabc 50.9 b c 49 4abc 53.5 C 45.6 a 45.8 a 50.2abc 51.9 b c
.67d .61bc .64 c d .53a .67d .63bcd .63 b c d .59b
± 1.6
.02
Basal 5% arenaceous flour? 5% gum arabicg 10% gum arabicg 5% xylose" 10% xylose" 5% lignin' 10% lignin1
SEMJ
a,b,c,d Means in the same column bearing different superscripts differ (P<.05). f
Means of triplicate groups of 5 chicks during the period 8 to 22 days posthatching. Described in Table 5.
^Described in Table 4. Described in Table 4. 'Described in Table 4. ••Pooled standard error of the mean.
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Treatment
Gain (g/day)
Basal 5% arenaceous flour^ 5% gum arabicS 10% gum arabicg 5% xylose" 10% xylose" 5% lignini 10% lignini
± 1.4
70.5a 72.9ab 81.0de 84.4 e 77_ 4 cd 78.4cd 76.3bc 76.3bc
82.lc 78.9 b c 65.4ab 70.4abc 71.3abc 83.5 C 60.7a 60.8 a
5.0
(Ac)
(/umoles/ml)
CecuiTl
'Described in Table 5.
Pooled standard error of the mean.
•Not detectable.
J
'Described in Table 4.
Described in Table 4.
"Described in Table 4.
1.3
20.7C 19.0 C 13.2 b 8.9 a 14.5 b 14.1b 14.7 b 13.4 b
8.8bc g jabc 5.9 a 6.6ab 8.0abc 7.4 a b 7.2ab 10.4 C .8
(Bu)
(Pr)
Molar %
a.b.^d.e,Means in the same column bearing different superscripts differ (P<.05).
SEMJ
1. 2. 3. 4. 5. 6. 7. 8.
Treatment
VFA con-
5.9
21. l a 23.3 a 43.5b 63.5cd 58.3bc 79.9 d 24.6 a 25.6 a
VFA concentration (Mmoles/ml)
TABLE 7. Total concentrations (iimoles/ml) and molar percentages of VFA s in cecal and l of chicks in Experiment 4
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FIBER EFFECTS ON CHICKS
DISCUSSION
Although fiber particle size did not influence (P<.05) performance of chicks fed semipurified diets, those fed the intermediate size particles (300 to 600 /xm) tended to have the highest rates of gain and feed efficiencies. This effect was noted with both bulking agents (PE and ACW), suggesting that particle size may be an important property of a fibrous ingredient, regardless of whether the fiber is potentially fermentable (ACW) or not (PE). Scott and Forbes (1958) noted that dietary fiber improved performance of chicks by simply narrowing the protein: calorie ratio, thus causing the chick to eat more of the most limiting amino acids to meet its energy needs. Results obtained with solka floe and PE-fed (300 to 600 urn and 600 to 1200 Urn) chicks in our studies support this work. Chicks fed ACW, however, performed better than those fed either solka floe or PE, indicating that possibly some component of ACW or microbial fermentation of the structural carbohydrates present in ACW may have resulted in beneficial effects on performance of the chick. This hypothesis is supported by data collected
in Experiment 2 (Table 4) where it was noted that certain of the components of ACW resulted in improvements in growth rates of chicks. However, chicks fed pectin, especially at the 8% level, performed poorly, as evidenced by low growth rates and feed efficiencies. Pectin-fed chicks ingested more (P<.05) feed than those fed other fiber types and excreted feces with a sticky consistency. The latter observation is supported by work of Wagner and Thomas (1977). These workers suggested that the growth depression seen with pectin was due to the growth of adverse intestinal microorganisms. In a subsequent paper, Wagner and Thomas (1978) suggested that a penicillinsensitive, spore-forming organism, which produced large amounts of gas and butyric acid fermentatively, depressed growth of chicks fed pectin. In certain instances, the feeding of D-xylose and L-arabinose, two primary components of hemicellulose, resulted in depressed growth rates of chicks (Wagh and Waibel, 1966; Baker, 1977), which may be attributed to direct competition with glucose for absorption sites (Ross et al, 1972) or poor metabolizability of these sugars (Wagh and Waibel, 1967). Wagh and Waibel (1966) found no significant differences between treatments when 10% xylosefed chicks were compared with controls. However, Baker (1977) found significant depressions at higher levels of xylose feeding. Very little work has been conducted on the nutritional value of the purified Kraft wood lignin used in this work. Most research to date has centered on its adsorptive properties (Kay et al, 1979; Rubio et al, 1979). Feeding both 4 and 8% lignin to chicks resulted in higher (P<.05) growth rates compared to control-fed chicks. Increased feed intake is a possible causal factor in the response, but it has recently been shown by Zemek et al. (1979) that certain phenolic constituents of lignin have antibioticlike properties. Also, Miller et al. (1979) found that various fractions isolated from wood hemicellulose extract, a chemically complex material high in phenolics, resulted in improvements in performance of chicks when incorporated into practical corn-soybean meal diets. Further evidence that phenolic compounds are involved in the growth-promoting properties of plant extracts can be noted in the work of Johanning and O'Dell (1981). They found that the cell wall material from cabbage and alfalfa which promoted guinea pig performance inhibited
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5 or 10% lignin was fed to chicks. Molar percent acetate was highest in ceca of chicks fed either level of gum arabic and lowest for basal and arenaceous flour-fed chicks. Chicks fed 10% lignin had the highest molar percent propionate (10.4), while those fed 5% gum arabic had the lowest (5.9). Molar percent butyrate was highest in ceca of chicks fed basal and 5% arenaceous flour diets. In the large intestine of chicks, VFA concentrations were low for basal, 5% arenaceous flour, and 5 and 10% lignin-fed chicks. The highest concentrations of VFA in the large intestine occurred when chicks were fed diets containing 10% gum arabic and 10% xylose. Level effects were noted for gum arabic and xylose. Molar percentages of acetate were high while molar percentages of propionate and butyrate were low. Ten percent gum arabic and 5 and 10% xylose-fed chicks had nondetectable levels of propionate. The highest molar percent propionate was noted for 10% lignin-fed chicks. Molar percent butyrate was highest in large intestinal contents of 5 and 10% gum arabicfed chicks and lowest in intestinal contents of chicks fed 10% xylose. No fiber level effect on molar percent was noted.
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Chicks started on cell wall carbohydratesupplemented diets at day 8 posthatching performed differently than those started at day 1. Chicks fed all treatment diets performed equivalent to or poorer than controls. A possible explanation for the different responses due to age may be that Hubbard chicks started at 8 days of age ingest such great quantities of feed (40 to 50 g/day) that rate of passage is increased to the extent that gut microbes do not have time to effectively ferment cell wall carbohydrate arriving at the ceca and large intestine. Adverse intestinal organisms may then compete with the normal flora, resulting in poorer animal performance. This is analogous to the situation which occurs with pectin-fed chicks (Wagner and Thomas, 1977).
Volatile fatty acid data suggested that different cell wall constituents affected gut microbial metabolism. Cecal VFA concentrations were highest for chicks fed basal and 10% xylose diets and lowest for chicks fed lignincontaining diets. Chicks fed 10% lignin had the highest molar percent cecal propionate, indicating that the cecal microbes of chicks fed this fiber source had a different fermentation pattern than those fed the other diets. Since phenolic components of lignin have been shown to have antibiotic-like properties (Zemek et al., 1979), this fermentation pattern may be of significance to the avian species. Much lower VFA concentrations (compared with cecal concentrations) were noted in the large intestine of chicks fed control diets and lignin-containing diets. Chicks fed 10% gum arabic and 10% xylose had the highest concentrations of large intestinal VFA's. Acetate was present in the greatest molar percent while molar percent butyrate was variable among treatments. Propionate was present in the lowest concentration. Chicks fed 10% lignin had the lowest molar percent acetate and the highest molar percent propionate, indicating a similar fermentation pattern to that of the ceca. In conclusion, it appears that cell wall constituents can have striking effects on avian performance and lower gut VFA patterns. More research is needed to define the chemical and physical properties of complex carbohydrates which will result in maximum performance of poultry. REFERENCES Annison, E. T., K. J. Hill, and R. Kenworthy, 1968. Volatile fatty acids in the digestive tract of the fowl. Br. J. Nutr. 22:207-216. Baker, D. H., 1977. Xylose and xylan utilization by the chick. Poultry Sci. 56:2105-2107. Bayer, R. C, W. H. Hoover, and F. V. Muir, 1978. Dietary fiber and meal feeding influence on broiler growth and crop fermentation. Poultry Sci. 57:1456-1459. Bryant, M. P., 1974. Nutritional features and ecology of predominant anaerobic bacteria of the intestinal tract. Am. J. Clin. Nutr. 27:1313-1319. Carmer, S. G., and M. R. Swanson, 1973. An evaluation of ten pairwise multiple comparisons procedures by Monte Carlo methods. J. Am. Stat. Assoc. 68:66-74. Erwin, E. S., G. J. Marco, and E. M. Emery, 1961. Volatile fatty acid analysis of blood and rumen fluid by gas chromatography. J. Dairy Sci. 44: 1768-1771. Goering, H. K., and P. J. Van Soest, 1970. Forage fiber analysis. ARS, USDA Handbook 379. Agric. Res. Serv., Washington, DC.
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growth of an unidentified anaerobic bacteria from guinea pig ceca. Extraction of the cell wall material with alkali resulted in the loss of pcoumaric and caffeic acids, two phenolics common to many plants (Hermann, 1980), and eliminated the antibacterial activity of the cell walls. Both of these acids are components of lignin. The best performances of 1-day-old chicks (Experiment 3) occurred when xylose and lignin were fed, suggesting that chicks fed these carbohydrates evidently establish a favorable gastrointestinal tract environment for utilization of the diet. Since a gut microbial population is established early in life, it appears that feeding certain cell wall constituents such as xylose and ball milled lignin from day 1 of age may result in an altered fermentation by gut microbes as evidenced by changes in the quantity and proportion of volatile fatty acids. Another possible explanation for the improvement in performance due to xylose supplementation may be that the specific activities of the enzymes catalyzing the reactions in the liver where xylose is converted to glucose are increased and thus xylose is being used as an energy source. The former explanation is probably more realistic, however, since even if lignin was degraded to its phenolic constituents and subsequently absorbed, no energy contribution to the animal would be possible. Also, since gum arabic was not well utilized by 1-dayold chicks (as evidenced by poor growth rates and feed efficiencies), it appears that the improvements due to the addition of cell wall carbohydrates is caused by a change in a microbial metabolism in the GI tract rather than a change in postabsorptive (tissue) metabolism.
FIBER EFFECTS ON CHICKS
2419. Rubio, M. A., S. Ingemar Falkehag, B. A. Pethica, and P. Zuman, 1979. The interactions of carcinogens and co-carcinogens with lignin and other components of dietary fiber. Pages 251—271 in Dietary fibers: Chemistry and nutrition. G. E. Inglett and S. Ingemar Falkehag, ed. Academic Press, New York, NY. Scott, H. M., and R. M. Forbes, 1958. The arginine requirement of chicks in relation to diet composition. Poultry Sci. 37:1347-1349. Thornburn, C. C , and J. S. Willcox, 1965. The caeca of the domestic fowl and digestion of the crude fiber complex. II. Experiments in vivo with fistulated birds, and the artificial and isolated caecum in vitro. Br. Poultry Sci. 6:33—43. Wagh, P. V., and P. E. Waibel, 1966. Metabolizability and nutritional implications of L-arabinose and D-xylose for chicks. J. Nutr. 9 0 : 2 0 7 - 2 1 1 . Wagh, P. V., and P. E. Waibel, 1967. Metabolism of L-arabinose and D-xylose by chicks. J. Nutr. 92: 491-496. Wagner, D. D., and O. P. Thomas, 1977. A rye type growth depression of chicks fed pectin. Poultry Sci. 56:615-619. Wagner, D. D., and O. P. Thomas, 1978. Influence of diets containing rye or pectin on the intestinal flora of chicks. Poultry Sci. 57:971-975. Zemek, J., B. Kosikova, J. Augustin, and D. Joniak, 1979. Antibiotic properties of lignin components. Folia Microbiol. 24:483-486.
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Heller, S. N„ L. R. Hackler, J. M. Rivers, P. J. Van Soest, D. A. Roe, B. A. Lewis, and J. B. Robertson, 1980. Dietary fiber: the effect of particle size of wheat bran on colonic function of young adult men. Am..J. Clin. Nutr. 33:1734-1744. Hermann, K., 1980. Pflanzenphenole als Inhaltsstoffe von Lebensmitteln Ernaehr. Umsch. 27: 75-80. Hungate, R. E., 1966. The Rumen and Its Microbes. Academic Press, New York, NY. Johanning, G. L., and B. L. O'Dell, 1981. Inhibition of a cecal anaerobe by a dietary fiber component. Fed. Proc. 40:485. (Abstr.) Kass, M. L., P. J. Van Soest, and W. G. Pond. 1980. Utilization of dietary fiber from alfalfa by growing swine. II. Volatile fatty acid concentrations in and disappearance from the gastrointestinal tract. J. Anim. Sci. 50:192-197. Kay, R. M., S. M. Strasberg, C. N. Petrunka, and M. Wayman, 1979. Differential absorption of bile acids by lignins. Pages 57—65 in Dietary Fibers: Chemistry and Nutrition. G. E. Inglett and S. Ingemar Falkenhag, ed. Academic Press, New York, NY. Miller, B. L., G. C. Fahey, Jr., and S. L. Schussler, 1979. Isolation of biologically active fractions from wood hemicellulose extracts. J. Anim. Sci. 48:1129-1134. Ross, A., A. W. Rubin, and J. J. Deren, 1972. Differential permeability of the proximal and distal rabbit small bowel. J. Clin. Invest. 51:2414—
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