Professional Documents
Culture Documents
S0308-8146(16)30301-6
http://dx.doi.org/10.1016/j.foodchem.2016.02.133
FOCH 18854
To appear in:
Food Chemistry
Received Date:
Revised Date:
Accepted Date:
7 August 2015
4 February 2016
21 February 2016
Please cite this article as: Li, X-k., Wang, J-z., Wang, C-q., Zhang, C-h., Li, X., Tang, C-h., Wei, X-l., Effect of
dietary phosphorus levels on meat quality and lipid metabolism in broiler chickens, Food Chemistry (2016), doi:
http://dx.doi.org/10.1016/j.foodchem.2016.02.133
This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers
we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and
review of the resulting proof before it is published in its final form. Please note that during the production process
errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
broiler chickens
Xue-ke Li a, b,1, Jin-zhi Wang a,1, Chun-qing Wang a, Chun-hui Zhang a*, Xia Li a,
5
6
10
11
* Corresponding author
12
13
14
15
Address: No. 2 Yuan Ming Yuan West Road, Haidian District, Beijing, China, 100193
16
17
18
19
20
21
22
Abstract
23
To analyze the influence of dietary phosphorus (P) levels on meat quality and lipid
24
25
level groups of H1 and H2, respectively) using 100 one-day-old broilers was
26
conducted. Results demonstrated that the quality of broiler chicken meat in deficient
27
or high P groups decreased relative to the normal group. High P diets resulted in
28
increased lightness, redness values, shear forces and decreased fatty acid contents and
29
intramuscular fat content in breast meat (p < 0.01). Compared with normal group,
30
lower malic enzyme activity, higher fatty acid synthase and AMP-activated protein
31
kinase activities were observed in the treatment groups (p < 0.05). Chickens fed with
32
normal diets had the lowest serum total cholesterol and triglyceride levels which
33
differed from that of other treatments (p < 0.05). High-P diets significantly decreased
34
the lipid accumulation in the liver (p < 0.01), whereas phosphorus levels in breast
35
meat increased significantly (p < 0.01). It can be concluded that deficient or higher P
36
levels could affect meat quality and expression of indicators on lipid metabolism of
37
broiler chickens.
38
39
40
1. Introduction
41
Chicken meat is considered a better choice for health diets because it contains less
42
43
compared with meat from other species (Riovanto et al., 2012). Due to growing
44
concerns on the relationship between diet and health, consumers are increasingly
45
demanding high-quality meat products. Consequently, factors that affect meat quality
46
47
48
factors. Among factors, phosphorus (P) is a critical and expensive mineral in poultry
49
nutrition. This mineral is used in energy pathways and during the synthesis of cell
50
51
dietary sources (Weinera et al., 2001). Studies on effects of dietary P levels on animal
52
growth and bone development showed that high P intake negatively impacts calcium
53
metabolism and bone properties, whereas low P diets will limit the growth of the
54
55
56
revisions to the P recommendations for broiler chicken feed (Yan et al., 2011). In
57
addition, farmers tend to use feed with more P than the recommended level (Lopez et
58
al., 2004). By far, there are barely any reports on the relationship between dietary P
59
60
61
All enzymes in the body need P (Takeda et al., 2012), but few reports describing how
62
63
64
AMP-activated protein kinase (AMPK), lipase, and main lipogenic enzymes (fatty
65
acid synthase (FAS), and malic enzyme (ME), etc. Among those enzymes, FAS can
66
alter the rates of the biosynthesis and hydrolysis of fatty acids (Smith, Witkowski, &
67
Joshi, 2003), and AMPK is primarily a critical regulator for energy metabolism (Kim
68
et al., 2004), regulating numerous intracellular pathways such as fatty acid oxidation
69
(Jonathan, John, & Bruce, 2012), development of fat cells and the deposition of
70
71
72
hypothesized that dietary P level may change the enzyme activities in broiler chickens,
73
thus affecting meat quality through lipid metabolism, since fatty acid composition and
74
biosynthesis play a role during the regulation for IMF contents (Camerona et al.,
75
2000).
76
To investigate the effects of dietary P levels on the meat quality and lipid
77
78
broilers. The chemical composition, meat color, cooking loss, tenderness, pH values,
79
fatty acid profiles and IMF content were determined to evaluate the differences in
80
meat quality affected by dietary P levels. Besides, to help understand how dietary P
81
levels affect lipid metabolism in broiler chickens, the activities of enzymes including
82
FAS, ME and AMPK in breast meat and liver were determined, trying to provide a
83
foundation for understanding the relation of dietary P levels and meat quality.
84
85
86
One hundred one-day old female broilers (Arbor Acres) were randomly allocated to
87
4 groups with 25 birds for each treatment. All birds were housed in metallic pens of
88
identical size (1.4 m 0.7 m 0.4 m) with 5 chickens per pen and provided with 12 h
89
light and dark cycles. The feeding experiments were conducted in animal testing
90
91
Chinese Academy of Agricultural Sciences. The birds were allowed to consume food
92
93
The experiment was divided into four groups according to available phosphorus
94
(AP) concentration of the feed: D (P deficient group), N (normal diet group), H1 (high
95
P group with available P 0.86% and 0.82%), and H2 (high P group with available P
96
1.16% and 1.10%). The nutrient levels of N group were based on the NRC (1994)
97
recommendations, and chickens were fed by basal diet. And on this basis we set
98
deficient group (D) and adequate groups (H1 and H2) with different P content in the
99
diet. The dietary P levels were mainly obtained by adding potassium dihydrogen
100
phosphate (feed grade) to the feed. The birds were given a diet based on corn and
101
soybean meal, and the diet included both starter and grower phases. The formulation
102
and nutrient levels of the feed are presented in Table 1. Contents of Ca and TP were
103
measured according to the AOAC (1935, 1996), while the others were calculated
104
values.
105
All chickens were weighed on a weekly basis, in the morning between 7:00 a.m and
5
106
8:00 a.m before the feed is offered. Body weight gain (BWG), feed intake (FI) and
107
feed conversion ratio (FCR) were calculated at the end of growth phase.
108
At 42 days of age, blood samples were collected from veins under the wings in all
109
live chickens. The serum was separated via centrifugation at 2500 g and 4 C for 15
110
min. The chicken weights, feed intake and mortality are shown in Table 2. And all the
111
live chickens were slaughtered and eviscerated. The liver and pancreas were removed,
112
rinsed in 0.9 % NaCl and immediately stored in liquid nitrogen for biochemical
113
analysis. The chickens were divided after evisceration. The whole breasts of each
114
chicken were cut off, and all samples were stored at -80 C until further analysis.
115
116
117
Waring blender (7012G, Texas, USA). Each of the following chemical analyses
118
involved a single sample from each chicken and experimental replication was across
119
all chickens in a particular treatment. The basic chemical composition of the meat was
120
determined according to the methods of AOAC (2000). The moisture contents were
121
122
protein content was determined using the Kjeldahl method (AOAC, 928.08), and the
123
total ash content was determined by weight after heating the samples in muffle
124
125
About 0.5 ~ 1.5 g of each homogeneous meat sample from each chicken was
126
weighed into crucible before being dried for 1 ~ 2 h at 110 C. The rest of procedure
127
follows that used to determine the total ash content. Afterwards, 5.0 mL of HCl was
128
added to the cooled crucible, and all rinses were transferred though a filter into a
129
volumetric flask. The same procedure was used for the blank. The TP of the breast
130
meat was determined from the ash solution by measuring the yellow color developed
131
132
Purkinje, Beijing, China) (AOAC, 1995). The phosphorus contents were expressed as
133
P.
134
135
The frozen breast meat samples were thawed at 4 C for 24 h and cut into 1 cm 1
136
cm 4 cm along the direction of the muscle fibers. The shear force values were
137
measured on raw meat samples using a Warner Bratzler shear force device (TA-XT2i
138
Plus, probe HDP/BSW, Stable Micro System Ltd., UK) according to the method of
139
140
141
cooked for 30 min at 80 C. Afterwards, the samples were cooled at room temperature
142
143
Meat color attributes including lightness (L*), redness (a*) and yellowness (b*)
144
145
Konica Minolta Sensing Inc., Japan) standardized against a white calibration plate.
146
147
148
149
The serum total cholesterol (TC) and triglyceride (TG) levels were measured using
150
151
152
The liver lipid extractions were conducted according to the procedures developed
153
by Lin et al., (2013). Briefly, the liver lipids were extracted using chloroform and
154
methanol (2:1, v/v). The TG in the liver was measured using commercial kits with a
155
156
157
The IMF content was measured using the methods described by Folch et al., (1957).
158
The total lipids were extracted in triplicates from each homogenized breast meat
159
160
The fatty acid composition in meat was determined by gas chromatography. Fatty
161
acid methyl esters (FAME) were prepared by esterification with acetyl chloride in
162
163
chloride in methanol in a tube that was tightly capped after filling with nitrogen. The
164
tube was then shaken vigorously and treated in a water bath for 2 h at 80 C. The tube
165
was washed using 3.0 mL sodium carbonate solution three times, and the resultant
166
solution was then centrifuged at 5000 g for 5 min. The supernatant was transferred
8
167
168
Next, 1.0 mL of the diluted fatty acid methyl ester solution was analyzed with a
169
Varian 450 Gas Chromatograph (Varian Inc., USA) equipped with a flame ionization
170
detector (FID) and a split injector. The injector and detector in the machine were set to
171
240 and 260 C, respectively. The helium carrier gas flow rate was 1.0 mL per min.
172
The initial oven temperature was 190 C, which was held for 7.5 min before
173
increasing at 3 C per min to 220 C. The final temperature was held for 2.5 min. In
174
the injector, the split valve was set to 1:60. The fatty acid composition of the sample
175
was calculated using the peak areas and methyl ester standard mixtures. The average
176
amount of each fatty acid was used to calculate the sum of the saturated fatty acid
177
178
179
180
The homogenates of breast meat and liver tissue were obtained according to the
181
methods of Underwood et al., (2008) with minor modifications. Briefly, the samples
182
183
184
185
using the supernatant obtained after centrifuging the homogenate (12000 g for 5
186
min at 4 C), and the differences between various samples were evaluated. The AMPK
187
activity was measured using commercially available kits (CUSABIO Biotech Co., Ltd,
9
188
189
190
The lipase activities in the pancreas and liver were analyzed according to the
191
method described by Knarreborg et al., (2003) with slight modifications. The samples
192
193
using an Ultra Turrax (T25 basic IKA, Germany) in an ice bath before being
194
centrifuged at 12000 g for 15 min at 4C. The supernatant was stored at -20C, and
195
the lipase activity was analyzed within 24 h using ELISA lipase kit (CUSABIO
196
Biotech Co., Ltd, Wuhan, China). Ten microliter of the supernatant was used when
197
198
199
The lipogenic enzyme (FAS and ME) activities in the breast meat and liver were
200
201
Briefly, the samples (1.0 g) were homogenized in 3.0 mL of ice-cold extraction buffer
202
203
204
at 12000 g for 30 min at 4 C, the supernatant was used to measure the enzymatic
205
activities (Rosebrough et al., 2002). The ELISA kits (CUSABIO Biotech Co., Ltd,
206
207
The optical density (OD) was measured at 450 nm with a microtiter plate reader
208
(Multiskan FC, Thermo Scientific, Vienna, Austria) to assay the AMPK, lipase and
10
209
210
211
For all analyses, surviving birds, pooled for each treatment served as the
212
experimental replicates. The data were analyzed using SPSS statistical software
213
(Version 19.0). All samples were used for experimental analysis, and single
214
215
(one-way ANOVA by Duncan) was used to assess the differences between the
216
treatments. A p-value of < 0.05 indicated significance. The data were expressed as the
217
218
3. Results
219
220
The effect of feeding diet with different P levels on feed consumption, feed
221
conversion ratio (FCR) and growth performance of broiler chickens has been shown
222
in Table 2. There were significant differences in final body weight among different
223
dietary P levels (p < 0.001). The N group had the highest values. The daily weight
224
gain of chickens fed with H1 and H2 diets was lower than N group (p < 0.05). The
225
results indicated that the chickens reared on high P diets (H1 and H2) had lower feed
226
consumption when compared to that fed the normal P diet (p < 0.05).
227
The chemical composition of the breast meat from broiler chickens is presented in
228
Table 3. Total phosphorus (TP) contents in breast meat reached 749 ~ 955 mg/kg,
11
229
230
231
and ash contents were observed in the treatment groups compared with that of N
232
except H2 group. However, the moisture content of meat from poultry fed the highest
233
P feed (H2) was significantly higher than the other three treatments (p < 0.01).
234
The effects of the dietary P level on the cooking losses, shear forces, meat color and
235
pH values are summarized in Table 3. The pH values and the cooking loss were not
236
significantly different compared with the normal diet group. The shear force and meat
237
color of breast meat showed extremely significant differences (p < 0.01) between
238
different groups. The treatment groups exhibited higher lightness (L*), redness (a*)
239
240
241
242
higher in H1 and H2 groups than in N group (p < 0.05). The TG concentrations in the
243
liver samples of H2 were significantly lower than in other groups (p < 0.05).
244
The IMF content was an important indicator of meat quality. The IMF content of N
245
group was 0.81 % higher than that of the treatment groups. The changes in the fatty
246
acid composition of the breast meat are shown in Table 4. It can be seen that
247
increasing dietary P levels decreased the fatty acid profiles significantly (p < 0.01)
248
except for that of stearic acid (C18:0) (p = 0.16). Similarly, the SFA, MUFA, PUFA
249
and total fatty acids decreased significantly with increasing of dietary P level (p <
12
250
0.01).
251
252
The AMPK plays an important role during the development of fat cells and the
253
deposition of IMF (Hardie, 2007). The AMPK activities were strongly influenced by
254
the dietary P levels (Fig. 1). The AMPK activity in the breast meat and liver of N
255
group is the lowest and that of D group is the highest. This result is contrary to the
256
changes observed in the IMF content of the breast meat, which showed the IMF
257
content of N group was significantly higher than that of treatment groups. The
258
negative correlation between AMPK activity and IMF content observed in the present
259
study is consistent with reported, which demonstrated that low AMPK activity is
260
261
Therefore, the AMPK activity is probably negatively associated with the IMF content
262
of breast meat.
263
The changes in the lipase activity in the pancreas and liver in broiler chickens are
264
shown in Fig. 1. Pancreatic lipase plays a substantial role during the digestion of
265
lipids (Knarreborg et al., 2003). The activity of pancreatic lipase increased when the
266
dietary P levels increased. The N group showed the highest lipase activity in liver
267
268
The effect of the dietary P levels on the lipogenesis modulation capacity was
269
assessed by determining the main lipogenic enzymes activities (ME and FAS) in
270
broiler chickens (Fig. 2). It can be seen that ME activity in N group was the highest in
13
271
the breast meat and the liver. Although the ME activities was lower in the
272
experimental groups (D, H1 and H2) versus N group, the FAS activities increased
273
continuously with the dietary P levels in breast meat. Whereas D group displayed
274
275
4. Discussion
276
The effects of the dietary P levels on the meat quality and lipid metabolism were
277
investigated. The total P and nonphytate phosphorus requirements for broilers during
278
the first three weeks and subsequent growth are 0.85 % and 0.80 %, 0.45 % and
279
280
broiler industry are common phenomena. Our results indicated that the high dietary P
281
282
chickens fed the high P diet was relatively lower than N group. TP content of the
283
breast meat increased significantly when the dietary P levels increased (p < 0.001),
284
indicating that excessive P consumption can lead to P deposits in muscle tissue. This
285
is consistent with the results of Marounek et al., (2008), which also demonstrated that
286
when the P intake in poultry diets reach a certain concentration, P levels in muscle
287
288
The meat quality attributes including the meat color, pH value, cooking loss, and
289
tenderness (Joo et al., 2013). Our results showed that the meat quality and chemical
290
composition of breast meat were significantly affected when the dietary P inclusion
291
levels increased. The chickens fed on diet excessive in P content (H2 group) had
14
292
significant higher moisture and ash contents (p < 0.05) than other groups. In this study,
293
the broiler chickens that consumed the experimental diets displayed significant impact
294
on meat color and shear forces in breast meat. The shear force is an objective
295
measurement of the tenderness of the meat (Lonergan et al., 2003). A lower shear
296
force value indicates that the meat is tenderer. In the current study, the shear force
297
values of the breast meat increased with increasing of dietary P levels, and the shear
298
force of N group is close to what was reported by Wattanachant et al. (2004). Our
299
300
301
In addition, the contents and composition of IMF are also important meat quality
302
characteristics (Yang et al., 2009), and its content indirectly affects the sensory
303
properties of meat, for example, tenderness and flavor, together with nutritional
304
quality (Joo et al., 2013; Symeon, 2013). Our results showed that IMF contents in N
305
group were significantly higher than that of deficient and high P group (p < 0.01),
306
indicating IMF content can be affected by the dietary P levels. The fatty acid
307
composition and biosynthesis play a role during the regulation for IMF contents
308
(Camerona et al., 2000) and its profiles affect the nutritional and sensory
309
characteristics of broiler chicken meat. Interestingly, breast meat from chickens fed
310
with deficient P in diet had relatively high fatty acid content than other group, and
311
high P dietary are likely to influence the SFA and PUFA, primarily the long chain
312
313
The distribution and deposition of the IMF is affected by the activities of various
314
key enzymes including ME, FAS and AMPK, which are related to lipid metabolism.
315
Our results showed that ME activities in liver were lower in treatments than that of N
316
group (p< 0.05), indicating the ME activity was inhibited when the P levels were high,
317
and the degree of inhibition interacted with the IMF contents. Previous study showed
318
the changes in the activities of the lipogenic enzymes (FAS) can alter the rates of the
319
biosynthesis and hydrolysis of fatty acids (Smith, Witkowski, & Joshi, 2003).
320
However, our results showed that the FAS activity in breast meat increased
321
significantly with increasing of dietary P levels, and the FAS activity in the liver and
322
breast meat displayed opposite trends relative to their fatty acid compositions.
323
Therefore, lower body weight and decrease in the fatty acid levels observed in the
324
treatment groups (D, H1 and H2) might be partly attributed to the decreased energy
325
intake as a result of reduced feed consumption, and resulted fatty acid oxidation to
326
327
Besides, our results showed that the IMF content decreased when the AMPK
328
activity increased in the breast meat, which agree with reported (Hardie, 2007;
329
Underwood et al., 2008). This can be partly explained as fatty acid synthesis is one of
330
the pathways most sensitive to AMPK inhibition (Underwood et al., 2008; Hardie et
331
al., 2003), and the activated AMPK can inhibit lipogenesis through phosphorylation,
332
333
334
enzyme during the absorption of dietary TG (Yang et al., 2014). Our results showed
335
lipase activity significantly increased with increasing of dietary P level. Chickens fed
336
337
chickens fed on diet deficient in P content, because TG is broken down into blood to
338
supply energy for life activities, causing increasing serum TC and TG concentrations
339
340
group is significantly lower than that of the other groups (p < 0.01). It is reported that
341
lipogenesis, the export of lipids, and synthesis and oxidation of fatty acid are
342
primarily achieved in the liver in endogenous lipid metabolism (Griffin et al., 1992;
343
Kersten, 2011), which are crucial steps for controlling TG accumulation in liver (Yang
344
et al., 2010). Previous reports have indicated that improving the serum lipid profile,
345
specifically increasing TC and TG, and decreasing the liver lipid accumulation and
346
lipid absorption are the most effective at decreasing the liver disorders (Yang et al.,
347
2014). During our study, high dietary P levels were not observed inducing fat
348
deposition in the liver. The results of lipid metabolism affected by P content in diet in
349
broilers may be explained from two aspects. On one hand, the metabolism of
350
lipoproteins may be facilitated. On the other, higher AMPK activity may inhibit the
351
TG synthesis in the liver. The effects of dietary P level on those parameters related to
352
353
5. Conclusion
354
The present study investigated the effects of dietary P on meat quality and
17
355
indicators for lipid metabolism. Results showed that higher P level in diet for broiler
356
357
inclusion levels in broiler diets may negatively affect meat quality, resulting in
358
decrease on IMF contents and fatty acid of breast meat, increase shear force (less
359
tenderness) when compared with the normal group. Different P level in diet affect key
360
enzymes related to lipid metabolism. Deficient or high P groups had increased AMPK
361
activities in breast meat and the liver, FAS activities in liver and lowed lipase activity
362
in liver, ME activities in breast meat and the liver compared with normal P group.
363
Lipase activity in pancreas and FAS activity in breast meat increased with increasing
364
365
chicken meat by regulating lipid metabolism through fed diets with different P levels
366
are needed.
367
Acknowledgements
368
The authors would like to thank the Fund of Agro-scientific Research in the Public
369
370
References
371
Alvarez, M. J., Diez A., Lopez-Bote C., Gallego M., & Bautista, J. M. (2000).
372
373
374
AOAC (1935). Official methods of analysis. AOAC Official Method 935.13. Calcium
18
375
376
377
378
379
380
AOAC (2000). Official methods of analysis (17th ed.). Moisture content. 950.46.
381
382
383
Camerona, N. D., Enserb, M., Nuteb, G. R., Whittingtonb, F. M., Penmana, J. C.,
384
Fiskena, A. C., Perryb, A. M., & Wood, J. D. (2000). Genotype with nutrition
385
interaction on fatty acid composition of intramuscular fat and the relationship with
386
387
Collin, A., Swennen, Q., Skiba-Cassy, S., Buyse, J., Chartrin, P., Le Bihan-Duval, E.,
388
Crochet, S. Duclos, M.J., Joubert, R., Decuypere, E., & Tesseraud, S. (2009).
389
390
391
153, 171-177.
392
Folch, J., Lees, M., & Sloane-Stanley, H. (1957). A simple method for the isolation
393
and purification of total lipids from animal tissue. The Journal of Biological
394
395
Griffin, H. D., Guo, K., Windsor, D., & Butterwith, S. C. (1992). Adipose tissue
19
396
lipogenesis and fat deposition in leaner broiler chickens. The Journal of Nutrition,
397
122, 363-368.
398
399
400
Hardie, D. G., Scott J. W., Pan D. A., & Hudson E. R. (2003). Management of cellular
401
energy by the AMP-activated protein kinase system. FEBS Letters, 546 (1),
402
113-120.
403
Hill, S. R., Knowlton, K. F., Kebreab, E., France, J., & Hanigan, M. D. (2008). A
404
405
406
Jonathan, S. O., John, W. S., & Bruce, E. K. (2012). AMPK functions as an adenylate
407
408
125-132.
409
410
411
412
Joo, S. T., Kim G. D., Hwang Y. H., & Ryu Y. C. (2013). Control of fresh meat quality
through manipulation of muscle fiber characteristics. Meat Science, 95, 828-836.
Kersten, S. (2011). Mechanisms of nutritional and hormonal regulation of lipogenesis.
European Molecular Biology Organization Reports, 2, 282-286.
413
Kim, J., Solis, R. S., Arias, E. B., & Cartee, G. D. (2004). Postcontraction insulin
414
415
416
96, 575-583.
20
417
Knarreborg, A., Jensen, S. K., & Engberg, R. M. (2003). Pancreatic lipase activity as
418
influenced by unconjugated bile acids and pH, measured in vitro and in vivo.
419
420
Lin, Y. L., Chang, Y. Y., Yang, D. J., Tzang, B. S., & Chen, Y. C. (2013). Beneficial
421
effects of noni (Morinda citrifolia L.) juice on livers of high-fat dietary hamsters.
422
423
Lonergan, S. M., Deeb N.; Fedler C. A., & Lamont, S. J. (2003). Breast meat quality
424
425
Lopez, H., Kanitz, F. D., Moreira, V. R., Wiltbank, M. C., & Satter, L. D. (2004).
426
427
428
Marounek, M., Skivan, M., Dlouh, G., & Beov, N. (2008). Availability of
429
430
laying hens 20 and 47 weeks old. Animal Feed Science and Technology, 146,
431
353-359.
432
433
434
435
436
437
438
Riovanto, R., Marchi, M. D., Cassandro, M., & Penasa, M. (2012). Use of near
439
440
441
442
443
vascular calcification, a related decrease in bone mass and changes in the aortic
444
445
Rosebrough, R. W., Poch, S. M., Russell, B. A., & Richards, M. P. (2002). Dietary
446
447
448
Smith, S., Witkowski, A., & Joshi, A. K. (2003). Structural and functional
449
organization of the animal fatty acid synthase. Progress in lipid research, 42,
450
289-317.
451
452
453
454
455
Takeda, E., Hironori, Y., Hisami, YO., & Taketani, Y. (2012). Dietary phosphorus in
bone health and quality of life. Nutrition Reviews, 70, 311-321.
456
Underwood, K. R., Means, W. J., Zhu, M. J., Ford, S. P., Hess, B. W., & Du, M.
457
458
fat content in longissimus dorsi muscle of beef cattle. Meat Science, 79, 394-402.
22
459
460
Wattanachant, Benjakul, & Ledward. (2004). Composition, Color, and Texture of Thai
Indigenous and Broiler Chicken Muscles. Poultry Science, 123-128.
461
Weinera, M. L., Salminenb, W. F., Larsonc, P. R., Barterd, R. A., Kranetze, J. L., &
462
463
464
Yan, Q. X., Tang, S. X., Musibau, A. B., Han, X. F., Zhou, C. S., Wang, M., Sun, Z. H.,
465
466
467
468
Yang, K. T., Lin C., Liu C. W., & Chen Y. C. (2014). Effects of chicken-liver
469
470
148-156.
471
Yang, X. J., Zhuang J. Y., Rao K. Q., Li X., & Zhao R. Q. (2010). Effect of early feed
472
473
474
Yang, Y., Song, J., Fu, R., Sun, Y., & Wen, J. (2009). The Expression of Can and
475
476
23
477
Figure captions
478
479
activities in breast meat (a) and liver (b) of broiler chickens. Effect of dietary
480
phosphorus level on lipase activities in pancreas (c) and liver (d) of broiler chickens.
481
The data are given as means SD. Different letters on data bars within each tested
482
483
normal group; H1, high P group with available P 0.86% and 0.82%; H2, high P group
484
485
486
breast meat (a) and liver (b) of broiler chickens. Effect of dietary phosphorus level on
487
fatty acid synthase (FAS) activities in breast meat (c) and liver (d) of broiler chickens.
488
The data are given as means SD. Different letters on data bars within each tested
489
490
normal group; H1, high P group with available P 0.86% and 0.82%; H2, high P group
491
24
24
26
AMPK in meat
36
22
20
18
ab
16
14
28
24
c
20
16
12
12
H1
H2
80
lipase in pancreas
76
H1
H2
Treatment
Treatment
AMPK in liver
32
36
ab
lipase in liver
72
68
64
34
60
56
a
32
30
28
52
26
492
493
H1
H2
H1
Treatment
Treatment
Figure 1.
494
25
H2
36
b 44
ME in meat
ME in liver
40
ME activity (U/L)
ME activity (U/L)
32
28
24
20
16
36
32
28
24
12
20
H1
H2
FAS in meat
90
80
60
a
a
40
78
72
66
54
496
84
60
30
495
FAS in liver
90
70
50
H2
d 96
100
H1
Treatment
Treatment
H1
H2
H1
Treatment
Treatment
Figure 2.
26
H2
Table 1: The composition and nutrient levels of diets for broiler chickens (air dry basis)
Starter phase (1~21 d)
D
H1
H2
H1
H2
Ingredients, %
Corn
Soybean meal
Rapeseed meal
Cottonseed meal
Fish meal
Limestone
CaHPO4
KH2 PO4
NaCl
Premix2
Total
61.40
19.50
6.00
5.00
3.00
1.50
0.10
0.20
0.30
3.00
100
60.00
20.50
5.50
5.00
3.00
1.10
0.60
1.00
0.30
3.00
100
60.70
18.70
3.80
4.00
6.00
0.80
0.70
2.00
0.30
3.00
100
59.00
19.00
4.50
3.55
5.60
0.85
0.70
3.50
0.30
3.00
100
62.00
19.00
4.70
5.00
4.50
1.50
0
0
0.30
3.00
100
62.00
19.00
4.00
3.50
5.90
1.20
0.20
0.90
0.30
3.00
100
61.00
18.50
3.80
4.00
6.00
0.90
0.50
2.00
0.30
3.00
100
59.70
19.10
4.10
3.50
5.60
0.80
0.70
3.20
0.30
3.00
100
Nutrient levels3
ME/(MJ/kg)
CP %
Ca %
TP %
AP %
Lys %
Met %
Met + Cys %
12.55
20.20
0.90
0.55
0.28
1.11
0.34
0.68
12.45
20.31
0.90
0.85
0.56
1.12
0.34
0.68
12.45
20.28
0.92
1.15
0.86
1.16
0.36
0.68
12.22
20.08
0.92
1.47
1.16
1.15
0.36
0.68
12.65
20.42
0.91
0.51
0.26
1.15
0.35
0.69
12.64
20.30
0.91
0.78
0.52
1.17
0.36
0.69
12.47
20.22
0.90
1.10
0.82
1.17
0.36
0.69
12.30
20.01
0.91
1.40
1.10
1.16
0.35
0.68
1) D, P deficient group; N, P normal group;H1, high P group with available P 0.86% and 0.82%; H2, high P group with available P 1.16% and 1.10%. 2) The premix
provided the following per kg of diets: VA 15000 IU, VD3 2700 IU, VE 60 mg, VB1 3 mg, VB2 9 mg, VB6 3 mg, VB12 0.03 mg, nicotinic acid 60 mg, pantothenic
acid 15 mg, biotin 0.36 mg, choline chloride 600 mg, Fe 90 mg, Cu 12 mg, Zn 75 mg, Mn 90 mg, I 0.35 mg, Se 0.25 mg. 3) Ca and TP were measured values, while
27
28
p-value
H1
H2
Initial/g
41.523.65
41.892.39
42.692.08
42.143.30
0.637
Final/g
1411135.56b
159796.68a
108094.54c
88370.27d
<0.001
BWG/g
32.543.99a
37.235.10a
24.584.63b
20.143.11b
0.002
FI (g/bird per d)
56.017.28ab
64.2312.00a
45.476.61bc
39.884.21c
0.012
FCR (g/g)
1.740.13b
1.720.10b
1.850.06ab
1.980.08a
0.022
Mortality/%
12
12
16
Weights
D, P deficient group; N, P normal group; H1, high P group with available P 0.86% and 0.82%; H2, high
P group with available P 1.16% and 1.10%. BWG, body weight gain; FI, feed intake; FCR, feed
conversion ratio.
29
Table 3: Effect of dietary P level on chemical composition and lipid metabolism in broiler chickens
Items
Moisture/ %
Crude
protein/ %
Ash/ %
Total
phosphorus
(mg/kg)
Shear force
(N)
Cooking
loss (%)
L*
a*
b*
pH value
lipid
metabolism
Serum TC
(mol/L)
Serum TG
(mmol/L)
TG in Liver
(mmol/L)
IMF in
muscle/ %
Treatments
p-value
H1
H2
73.590.30a
73.740.36a
73.770.39a
75.130.37b
<0.001
24.40.95
24.150.44
23.840.79
23.451.23
0.457
1.590.20ab
1.710.13a
1.680.14a
1.340.19b
0.037
74967a
87637b
90735bc
95523c
<0.001
11.430.70b
8.950.72c
10.591.02b
12.920.64a
<0.001
22.950.74
23.191.06
23.510.60
24.531.87
0.312
53.562.21ab
1.410.48ab
4.190.85a
5.730.13
51.871.72a
1.040.29a
5.390.96b
5.840.12
52.621.38a
1.860.51b
4.770.99ab
5.760.13
55.311.55b
2.430.17c
5.801.31b
5.730.08
0.009
0.001
0.032
0.205
3.390.31ab
3.200.15a
3.560.33b
3.680.34b
0.041
0.370.04ab
0.340.03a
0.440.06bc
0.460.07c
0.009
3.390.29a
3.560.05a
3.480.14a
2.550.39b
0.004
0.620.02a
0.810.03c
0.700.01b
0.610.01a
<0.001
D, P deficient group; N, P normal group; H1, high P group with available P 0.86% and 0.82%; H2, high
P group with available P 1.16% and 1.10%. Results are represented as mean SD. Dierent letters in
the same row indicate significant dierences (p < 0.05). L*, Lightness; a*, Redness; b*, Yellowness.
TC= Total cholesterol; TG= Triglyceride; IMF= Intramuscular fat contents.
30
Table 4: Effect of dietary P level on fatty acids profile of breast meat in broiler chickens
Treatments
Items
p-value
H1
H2
C16:0
141.659.22a
123.994.36b
123.454.53b
95.07.35c
<0.001
C16:1
15.410.17a
13.941.43a
10.580.04b
7.481.21c
<0.001
C18:0
83.492.14
76.423.97
75.622.71
74.967.48
0.160
C18:1
121.653.31a
92.548.94c
104.024.86b
55.912.26d
<0.001
C18:2
132.888.09a
108.935.84c
120.335.98b
91.33.29d
<0.001
C18:3
5.00.21a
3.950.56bc
4.430.45ab
3.190.34c
0.004
SFA
225.1411.36a
200.418.33b
199.077.24b
169.9614.83c
0.002
MUFA
137.063.48a
121.8510.37b
114.64.9b
63.393.47c
<0.001
PUFA
137.888.3a
112.229.81c
124.768.69b
94.483.63d
<0.001
Total fatty
acids
500.0823.14a
419.1125.97b
438.435.31b
327.8319.35c
<0.001
D, P deficient group; N, P normal group; H1, high P group with available P 0.86% and 0.82%; H2,
high P group with available P 1.16% and 1.10%. Results are represented as mean SD and
expressed as mg FA/100 g fresh meat. Dierent letters in the same row indicate significant
dierences (p < 0.05). SFA= saturated fatty acid; MUFA= mono-unsaturated fatty acid; PUFA=
poly-unsaturated fatty acid.
31
32