谷氨酰胺和γ-氨基丁酸对肉鸡抗热应激和肉品质的影响及机理探讨
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摘要
本文通过体内试验研究了循环热应激(30-34℃)对肉鸡生长性能、屠宰性能、胴体性状、血清生化指标、肌肉品质、骨骼肌中G1n和GABA代谢、HSP70、AMPK信号通路相关指标的影响及外源性Gln和GABA的添加效果和互作效应,探讨了可能存在的机理,分析了骨骼肌部分指标与肌肉品质间的相关性。并进一步通过体外试验探讨了热应激条件下家禽骨骼肌细胞HSP70、AMPK问的调控关系及外源性Gln、GABA的干预作用,为氨基酸用于家禽抗热应激提供理论基础。
1.体内试验：按照2因素2水平(2×2)加正对照组(PC组)的实验设计进行。选择健康的10日龄(d)健康AA公鸡360只,在适温条件下,常规饲养至21d时随机分为5组(每组6重复,每重复12只)。其中,四个热应激组(2×2)在22-42d进行循环热应激处理(每日9：00-14：00,温度从30℃升至34℃,14：00-18：00温度从34℃降至30℃,其他时间平均23℃左右),在基础日粮中分别添加Gln(0、5g/kg)和GABA (0、100mg/kg)。PC组在适温条件饲养(约23℃),饲喂基础日粮。于28、35、42d进行生长性能分析,并分别从各重复组抽取3只接近平均体重的样鸡用于屠宰性能、胴体品质、血清生化指标、肌肉品质、骨骼肌中Gin和GABA代谢、HSP70、AMPK信号通路相关组分的分析,对部分指标与肌肉品质之间的相关性进行分析,结果如下：
1.1循环热应激降低了22-42d肉鸡增重(WG)和采食量(FI),42d时的屠体重(CW)、半净膛重(HEW)、全净膛重(EW)、胸肌重(BMW)、胸肌率(BMY)、腿肌重(LMW)、腹脂重(AFW)、腹脂率(AFY)、肌间脂肪宽(IFW)、皮下脂肪厚(SFT)；增加了肉鸡料重比(F/G)和腿肌率(LMY)。日粮添加5g/kg Gln增加了22-42d热应激肉鸡WG、FI、CW、HEW、EW、BMW、BMY、LMW、AFW、AFY、 IFW、SFT;降低了F/G,对LMY有降低趋势。日粮添加100mg/kg GABA增加了22-42d热应激肉鸡WG、FI、EW、BMW、BMY、AFW、AFY、SFT;对CW、HEW、IFW有增加趋势。两种氨基酸在影响循环热应激肉鸡WG、FI、F/G、EW、EY、BMW、 BMY、AFW、AFY、IFW和SFT方面存在协同效应。在影响生产性能方面,Gln的作用具有较好的持续性,GABA在第1周时作用较强,随后明显减弱。
1.2循环热应激降低了28d、35d、42d肉鸡血清TP、葡萄糖、甘油三酯、Gin、Glu、T4、Ins水平、ALP活性,22d、35d血清GABA水平；增加了22d、35d、42d肉鸡血清CS、GN水平、GOT、GPT、LDH、CK、NOS活性。日粮添加5g/kg Gln增加了22d、35d、42d热应激肉鸡血清TP、Gln、Ins水平,降低了血清甘油三酯、CS、GN水平、GOT、CK、NOS活性。日粮添加100mg/kg GABA增加了22d、35d、42d热应激肉鸡血清TP、Gln、GABA、T3水平、ALP活性,对甘油三酯、Gln、T4、 CS、Ins、GN水平、GOT、GPT、LDH、CK、NOS活性的影响存在明显的时间特异性。两种氨基酸在影响循环热应激肉鸡血清TP、甘油三酯、GABA、Gln、T3、T4、 Ins、GN水平、ALP、LDH和CK活性方面存在明显的协同效应,在影响转氨酶(GOT、 GPT)和NOS方面仅在第1周时较显著。
1.3循环热应激增加了28、35、42d肉鸡胸肌pH、L*、水分、CP、CF、CA、 TL、F、MFDS,腿肌pH、L*、CA, DL、SF、MFDS;降低了肉鸡胸肌DL、CLMFDM,腿肌a*、b*、CP、CF、CL、TL、CL、MFDM。日粮添加5g/kg Gln降低了胸肌L*、SF、MFDS,腿肌L*、DL、SF、MFDS,增加了胸肌CP、CF、CA、TL、 DL、MFDM,腿肌a*、CP、CF、水分、CL、MFDM.日粮添加100mg/kg GABA降低了胸肌pH、TL、SF、腿肌pH、DL、SF,增加了胸肌a*、水分、CP、CF,腿肌a*、CF。两种氨基酸在影响循环热应激肉鸡胸肌pH、b*、水分、CP、WHC、TL、SF,腿肌pH、L*、a*、水分、CP、WHC、TL、SF方面存在明显的协同效应。此外,通过日粮Gln和GABA可以提高肌肉在冷藏过程中的色泽稳定性。
1.4循环热应激降低了28d、35d、42d肉鸡胸肌、腿肌的Gln、Glu、GABA含量、Glnase、GAD活性,增加了胸肌、腿肌的GS、GABA-T活性。日粮添加5g/kg Gln增加了胸肌的Gln、Glu含量、Glnase、GAD活性,腿肌的Gln、Glu、GABA含量、GAD活性；降低了胸肌、腿肌的GS活性,对胸肌的GABA含量、GABA-T活性、腿肌的Glnase、GABA-T活性有不同程度的影响。日粮添加100mg/kg GABA增加了胸肌的Gln、Glu含量、Glnase、GAD活性,腿肌Gln含量、GAD活性；降低了胸肌的GABA-T活性、腿肌的GS活性。两种氨基酸对循环热应激肉鸡胸肌、腿肌的Gln、 GABA含量、Glnase、GS、GABA-T活性的影响存在明显的协同效应。42d肉鸡胸肌、腿肌的Gln、GABA含量、Glnase、GS活性与L*、CP、CF、CL、MFDS和MFDM间存在显著的相关性。
1.5循环热应激降低了28d、35d、42d肉鸡胸肌、腿肌糖原、葡萄糖、乳酸含量,增加了胸肌、腿肌的AMP/ATP、AMPK含量、AMPK活性、HSP70含量、AMPKa2mRNA、HSP70mRNA表达量。日粮添加5g/kg Gln增加了胸肌、腿肌的HSP70含量、HSP70mRNA表达量、胸肌的葡萄糖、腿肌的糖原；降低了胸肌、腿肌的AMP/ATP、 AMPK含量、AMPK活性。日粮添加100mg/kg GABA增加胸肌的糖原、葡萄糖、腿肌的葡萄糖、乳酸,降低了胸肌、腿肌的AMP/ATP、AMPK活性、腿肌的HSP70含量、HSP70mRNA表达量。两种氨基酸对循环热应激肉鸡胸肌的葡萄糖、乳酸、腿肌的糖原、乳酸、胸肌和腿肌的AMP/ATP、AMPK活性、HSP70mRNA表达量存在明显的协同效应。42d肉鸡胸肌和腿肌的糖原、葡萄糖、乳酸与水分、CP、CF, AMP/ATP. AMPK含量、AMPK活性与L*、水分、CP、CF、WHC、DL、CL、MFDS和MFDM间,HSP70含量与WHC、TL、DL间存在较为显著的相关性。
2.体外试验：研究热应激影响家禽骨骼肌细胞HSP70、AMPK的调控关系及外源性Gln. GABA的干预作用。以1-2×106个/mL密度的成肌细胞进行试验。处理后置于37℃水浴中进行3h修复,收集细胞供分析。
2.1热应激条件下HSP70表达与AMPK活性变化时差性分析
共5组,每组6重复,基础培养基培养3h后进行热应激处理(42℃和45℃,0-120min)。结果表明：热应激提高了细胞的HSP70表达量和AMPK活性,以45℃处理的提高速度更快,程度更大,且以AMPK活性的变化速度快于HSP70表达量的变化。
2.2HSP70表达与AMPK活性问的调控关系分析
(1)QD抑制HSP70表达试验：共7组,每组6重复。用添加浓度为0-50umol/L的QD的基础培养基预培养3h,进行42℃和45℃、30min的热应激处理。结果表明：QD对HSP70表达的抑制作用存在明显的温度特异性,仅45℃时表现显著。同时,45℃时的AMPK活性具有不同程度的增加,显示HSP70能在一定程度上抑制/AMPK活性。
(2)AICAR激活AMPK试验：共7组,每组6重复。用添加浓度为0-0.5mmol/L的AICAR的基础培养基培养3h处理。结果表明：AICAR能有效激活适温条件下的骨骼肌细胞AMPK活性,但对HSP70的表达量无影响。
2.3Gln、GABA对细胞HSP70与AMPK的影响
各取5组,每组6重复。在基础培养基中分别添加0-20mmol/L Gln和0-0.8mmol/L GABA预培3h后,进行45℃,30mmin热应激处理。结果表明：Gln能有效增加热应激条件下骨骼肌细胞的HSP70表达量,同时抑制AMPK活性的增加。GABA能在一定程度上抑制AMPK活性的增加,但对HSP70表达量无影响。
2.4Gln、GABA对极端热应激骨骼肌细胞成活率的影响
各取3组,每组6重复。分别用添加0、10、20mmol/L Gln和0、0.4、0.8mmol/L GABA的基础培养基预孵3h后,采用48℃高温处理3h、6h,分析细胞成活率。结果表明：添加10、20mmol/L Gln提高了热应激处理3h、6h后的骨骼肌细胞成活率；添加0.4、0.8mmol/L GABA对细胞成活率无明显影响。
根据研究结果可以得出结论：
(1)本研究采用的30-34℃循环热应激对22-42d肉鸡生长性能、胴体性状、血清主要代谢物、激素和代谢酶等生化指标产生了显著的负面影响,也显示了机体对热应激的适应性。日粮添加5g/kg Gin、100mg/kg GABA能有效缓解循环热应激对上述指标带来的负面作用,同时表现出了明显的协同效应和时间特异性。
(2)从物质代谢角度分析,循环热应激显著影响了血清和骨骼肌中的Gln、Glu、 GABA水平,以及三种氨基酸间的代谢酶。通过日粮添加5g/kg Gln、100mg/kg GABA可以有效改善循环热应激条件下肉鸡骨骼肌中Gln、GABA的代谢状况,同时表现出了一定的协同效应和骨骼肌组织差异性。
(3)循环热应激明显造成了肉鸡机体的低能量状态,激活了骨骼肌中AMPK信号通路,促进了骨骼肌中HSP70转录和表达。通过日粮添加5g/kg Gln、100mg/kg GABA可以改善机体的能量状况,抑制骨骼肌AMPK信号通路,且表现出了一定的协同效应。同时说明,Gin的这种积极作用可能在一定程度上与促进HSP70表达有关,而GABA的改善作用与HSP70无关,可能与其神经系统和生理调控方面功能有关。
(4)循环热应激对胸肌、腿肌的化学组成和肉品质指标产生了显著的负面影响。通过日粮中添加5g/kg Gln、100mg/kg GABA对于缓解热应激对这些指标带来的负面影响具有积极作用,其中的机制与骨骼肌中Gin代谢变化、(?)AMPK信号通路激活状况、HSP70表达量的变化有密切关系,且表现出了一定的协同效应和骨骼肌组织差异性。
(5)通过体外骨骼肌细胞试验证明,热应激能激活肌细胞AMPK信号通路,促进HSP70的表达。热应激细胞中HSP70的表达可在一定程度上抑制AMPK信号通路的激活,起到保护细胞作用。在适温条件下细胞中AMPK的激活并不能导致HSP70表达量的变化。同时证明,外源性Gln、GABA均能抑制热应激骨骼肌细胞AMPK信号通路,但仅有Gln能有效保护热应激细胞,其作用与促进HSP70表达密切相关。
This experiment in vivo was conducted on broilers to evaluate the effects of circular heat stress (CHS) on growth performance, slaughter performance, carcass characteristic, serum biochemical parameter, meat quality, skeletal metabolism of Glutamine (Gln) and gamma-aminobutyric acid (GABA), heat shock protein (HSP70) and relative indexes in AMP-activated protein kinase (AMPK) signal path. The effects and interactions of dietary Gln and GABA on the broilers under CHS and the possible mechanism were also be evaluated. Moreover, the correlations between some skeletal indexes and meat quality indexes were analyzed. Another experiment in vitro was also conducted to evaluate regular relationship between HSP70and AMPK in poultry skeletal cell under heat stress (HS) and intervention of exogenous Gln and GABA on these indexes. Resolution of these issues will provide theoretical foundation for applying Gln and GABA against HS.
1. Experiment in vivo
The experiment was a2×2plus positive group (PC) of the treatments with two levels each of Gln (0and5g/kg Gln of diet) and GABA (0and100mg/kg GABA of diet). A total of36010-day-old healthy male Arbor Acres broiler chicks were procured and were housed from day10to day21in thermostatically controlled cages and were fed with same basic compound daily diet. On day22, after overnight (12h) feed withdrawal, chicks were wing-banded and randomly assigned to five treatment groups (6replicates of12birds per cage). The chicks of the four treatment groups (2×2) were housed in a circular high temperature controlled room (from9:00to14:00at30-34℃, then from14:00to18:00at34-30℃, and rest of the time at average23℃) while the PC group was housed in another optimal temperature controlled room (average23℃). Growth performance for each group was recorded from22to42days of age. At28.35,42days of age, three broilers from each cage with a body weight (BW) close to the group average was selected to analysis slaughter performance, carcass characteristics, serum biochemical parameters, meat quality, skeletal metabolism of Gin and GABA, HSP70and relative indexes in AMPK signal path, and to evaluate the correlations between some skeletal indexes and meat quality indexes. Results as below:
1.1CHS decreased weight gain (WG), feed intake (FI) of broiler from28d to42d old, carcass weight (CW), half-eviscerated weight (HEW), eviscerated weight (EW), breast muscle weight (BMW), breast muscle yield (BMY), leg muscle weight (LMW), abdominal fat weight (AFW), abdominal fat yield (AFY), intermuscle fat width (IFW)&subcutaneous fat thick (SFT) of42d old broiler, while increased feed-to-gain ratio (F/G)&leg muscle yield (LMY). The diet supplemented with5g/kg Gln increased WG, FI, CW, HEW, EW, BMW, BMY, LMW, AFW, AFY, IFW&SFT, while decreased F/G and had decreasing tendency for LMY of28-42d old broilers under CHS. The addition of100mg/kg GABA increased WG, FI, EW, BMW, BMY, AFW, AFY&SFT, while had increasing tendency for CW, HEW&IFW of28-42d old broilers under CHS. Different extent synergistic effects were found between Gln and GABA for WG, FI, F/G, EW, eviscerated yield (EY), BMW, BMY, AFW, AFY, IFW&SFT. With regard to the effects of Gln and GABA on growth performance,Gln had persistent action, while GABA had gradually weakened action.
1.2CHS decreased concentrations of TP, Glucose, triglyceride, Gln, Glu and T4, activities of ALP in serum of28,35,42d old broilers, and GABA content of28,35d old broilers, while increased concentrations of CS&GN, activities of GOT, GPT, LDH, CK&NOS in serum of28,35,42d old broilers. The diet supplemented with5g/kg Gln increased concentrations of TP, Gln&Ins, while decreased concentrations of triglyceride, CS&GN, activities of GOT, CK&NOS in serum of28,35,42d old broilers under CHS. The addition of100mg/kg GABA increased concentrations of TP, Gln, GABA&T3, ALP activities, while had obvious time-specifically effects on concentrations of triglyceride, Glu, T4, CS, Ins&GN, activities of GOT, GPT, LDH, CK&NOS in serum of28-42d old broilers under CHS. Different extent synergistic effects were found between Gln and GABA for serum concentrations of TP, triglyceride, GABA, Glu, T3, T4, Ins&GN, activities of ALP, LDH&CK, and GOT, GPT&NOS were found only in1st week under HS.
1.3CHS increased pH, L*, moisture, CP, CF, CA, TL, SF&muscle fiber density (MFDS) of breast muscle, pH, L*, CA, DL, SF&MFDS of leg muscle, while decreased DL, CL&muscle fiber diameter (MFDM) of breast muscle, a*, b*, CP, CF, CL, TL, CL&MFDM of leg muscle of28,35,42d old broilers. The diet supplemented with5g/kg Gln decreased L*, SF&MFDS of breast muscle, L*, DL, SF&MFDS of leg muscle, while increased CP, CF, CA, TL, DL&MFDM of breast muscle, a*, CP, CF, moisture CL&MFDM of leg muscle of broilers under CHS. The addition of100mg/kg GABA decreased pH, TL&SF breast muscle, pH, DL&SF of leg muscle, while increased a*, moisture, CP&CF of breast muscle, a*&CF of leg muscle of broilers under CHS. Different extent interactions were found between Gln and GABA for pH, b*, moisture, CP, WHC, TL&SF of breast muscle, pH, L*, a*, moisture, CP, WHC, TL&SF of leg muscle. Moreover, dietary Gln and GABA could improve color stability of breast and leg muscle during cold-storage.
1.4CHS decreased concentrations of Gln, Glu&GABA, activities of Glnase&GAD, while increased activities of GS&GABA-T in breast and leg muscle of28,35,42d old broilers. The diet supplemented with5g/kg Gin increased concentrations of Gln&Glu, activities of Glnase&GAD in breast muscle, concentrations of Gln, Glu&GABA, activities of GAD in leg muscle, while decreased GS activates of breast and leg muscle, and had different effects on GABA concentration and GABA-T activity in breast muscle, Glnase, GABA-T activities in leg muscle of broilers under CHS. The addition of100mg/kg GABA increased concentrations of Gln&Glu, activities of Glnase&GAD in breast muscle, Gln concentration and GAD activity in leg muscle, while decreased GABA-T activity in breast muscle and GS activity in leg muscle of broilers under CHS. Significant synergistic effects between dietary Gln and GABA on Gln content, GABA content, Glnase activity, GS activity&GABA-T activity were found. There were significant correlations among indexes of Gln content, GABA content, Glnase activity&GS activity and indexes of L*, CP, CF, CL, MFDS&MFDM in breast&leg muscle of42d broilers.
1.5CHS decreased concentrations of glycogen, Glucose&lactic acid, while increased AMP/ATP, AMPK content, AMPK activity, HSP70content, expression of AMPK a2mRNA and HSP70mRNA in breast and leg muscle of28,35,42d old broilers. The diet supplemented with5g/kg Gin increased HSP70content, HSP70mRNA expression in breast and leg muscle, glycogen content in breast muscle, Glucose content in leg muscle, while decreased AMP/ATP, AMPK content, AMPK activity in leg muscle of broilers under CHS. The addition of100mg/kg GABA increased glycogen and glucose contents in breast muscle, glucose and lactic acid contents in leg muscle, while decreased AMP/ATP and AMPK activity in breast and leg muscle, HSP70content, HSP70mRNA expression in leg muscle of broilers under CHS. Significant interactions between dietary Gln and GABA on glucose and lactic acid contents in breast muscle, glycogen and lactic acid in leg muscle, AMP/ATP, AMPK activity, HSP70mRNA expression in breast and leg muscle were found. There were significant correlations among indexes of glycogen, glucose&lactic acid and indexes of moisture, CP&CF, indexes of AMP/ATP, AMPK content&K activity and indexes of L*, moisture, CP, CF, WHC, DL, CL, MFDS&MFDM in breast&leg muscle. The significant correlations among HSP70and WHC, TL&DL were found.
2. Experiment in vitro
This experiment in vitro was conducted to evaluate the relationship between HSP70and AMPK in poultry skeletal cell under heat stress (HS), and the intervention of exogenous Gin and GAB A. The skeletal cell with density of1-2×106/ml were used in the experiments. After experimental treatment, the skeletal cell was repaired in37℃water bath for3h, and then was collected for analysis.
2.1Analysis on time difference of HSP70and AMPK under HS
A total of five treatment groups (6replicates of per group) were used in this experiment. HS (42℃and45℃,0-120min) was conducted on skeletal cell after culture in basic medium for3h. The results showed that HS increased HSP70expression and AMPK activity and the HS with45℃had more quickly and heavily increasing on the two indexes The increasing of AMPK activity was proceeding to HSP70expression.
2.2Analysis on regulation relationship between HSP70expression and AMPK activity
(1) Experiment about Quercetin (QD) inhibits HSP70expression:
A total of seven treatment groups (6replicates of per group) were used in this experiment. HS (42℃and45℃,30min) was conducted on skeletal cell after culture in basic medium with supplementation of0-50umol/L QD for3h. The results showed that inhibition of QD on HSP70expression showed obvious temperature-specificity, and the inhibition only displayed under45℃condition. Meanwhile, the increasing of AMPK activity under45℃showed that HSP70could inhibit AMPK activity to a certain extent.
(2) Experiment about AICAR activates AMPK:
A total of seven treatment groups (6replicates of per group) were used in this experiment. Skeletal cell was cultured in basic medium with supplementation of0-0.5mmol/L AICAT for3h under thermoneutrality. The results showed that AICAR could activate AMPK, but HSP70expression was not effected by AICAR.
2.3Effect of Gln and GABA on HSP70and AMPK
Two totals of five treatment groups (6replicates of per group) were used respectively in this experiment. HS (45℃,30min) was conducted on skeletal cell after cultured in basic medium supplemented with0-20mmol/L Gln or0-0.8mmol/L GABA for3h. The results showed that Gln could increase HSP70expression, while inhibit AMPK activity, and GABA could inhibit AMPK activity to a certain extent, but had no effect on HSP70expression in skeletal cell under HS.
2.4Effect of Gln and GABA on survival rate of skeletal cell under extremely HS
Two totals of three treatment groups (6replicates of per group) were used respectively in this experiment. HS (48℃,3h and6h) was conducted on skeletal cell after cultured in basic medium supplemented with0,10,20mmol/L Gln or0,0.4,0.8mmol/L GABA for3h. The results showed that10,20mmol/L Gln could increase survival rate of skeletal cell under HS for3h,6h (P<0.05or0.01), but0.4,0.8mmol/L GABA had no effect.
As stated above, the conclusions are as follows:
(1) The CHS (30-34℃) had obviously negative effects on growth performance, slaughter performance, carcass characteristics and serum biochemical parameters (main metabolites, hormones, metabolic enzymes), which also showed adaptability against HS. The diet supplemented with5g/kg Gln and100mg/kg GABA offered a potential nutritional strategy to prevent HS-related depression in these indexes of broiler, and also showed significant synergistic effect and time-specificity.
(2) From the material metabolism, CHS affected significantly Gln, Glu and GABA contents in serum and skeletal muscle and metabolic enzyme among them. Dietary Gln (5g/kg) and GABA (100mg/kg) improved metabolic state of Gln and GABA in skeletal muscle of broiler under CHS, and also showed a certain synergistic effect and skeletal tissue-variability.
(3) CHS obviously formed low-energy state of organism, activated AMPK signal path, and increased HSP70expression in skeletal muscle of broiler. Dietary Gln (5g/kg) and GABA (100mg/kg) improved energy state, inhibited AMPK signal path in skeletal muscle, and also showed a certain interactions between this two amino acids. The results also indicated that the positive action of Gln might be relevant to promoting HSP70expression. But the improvement of GABA was unrelated to HSP70expression, which may be involved in function of GABA in nervous system and physiological regulation.
(4) CHS had significant negative effects on chemical compositions and quality of breast and leg meat. Dietary Gln (5g/kg) and GABA (100mg/kg) offered a potential nutritional strategy to prevent HS-related depression in these indexes of broiler, and also showed a certain interaction and skeletal tissue-variability. The impossible mechanism of CHS affects meat quality is related to metabolic change of Gln, activated state of AMPK signal path, and variation of HSP70expression to agreater extent
(5) By test in vitro proved that HS activated AMPK signal path, and increased HSP70expression in skeletal muscle cell. Elevated HSP70expression could inhibit the activation of AMPK signal path in skeletal cell under HS and thus protect skeletal cell from HS-relative damage. The activation of AMPK could not affect HSP70expression in skeletal cell under thermoneutrality. Exogenous Gln and GABA could both inhibit activation of AMPK, but only Gln had effective protection to skeletal cell under HS, which was related to increasing HSP70expression.
1.体内试验：按照2因素2水平(2×2)加正对照组(PC组)的实验设计进行。选择健康的10日龄(d)健康AA公鸡360只,在适温条件下,常规饲养至21d时随机分为5组(每组6重复,每重复12只)。其中,四个热应激组(2×2)在22-42d进行循环热应激处理(每日9：00-14：00,温度从30℃升至34℃,14：00-18：00温度从34℃降至30℃,其他时间平均23℃左右),在基础日粮中分别添加Gln(0、5g/kg)和GABA (0、100mg/kg)。PC组在适温条件饲养(约23℃),饲喂基础日粮。于28、35、42d进行生长性能分析,并分别从各重复组抽取3只接近平均体重的样鸡用于屠宰性能、胴体品质、血清生化指标、肌肉品质、骨骼肌中Gin和GABA代谢、HSP70、AMPK信号通路相关组分的分析,对部分指标与肌肉品质之间的相关性进行分析,结果如下：
1.1循环热应激降低了22-42d肉鸡增重(WG)和采食量(FI),42d时的屠体重(CW)、半净膛重(HEW)、全净膛重(EW)、胸肌重(BMW)、胸肌率(BMY)、腿肌重(LMW)、腹脂重(AFW)、腹脂率(AFY)、肌间脂肪宽(IFW)、皮下脂肪厚(SFT)；增加了肉鸡料重比(F/G)和腿肌率(LMY)。日粮添加5g/kg Gln增加了22-42d热应激肉鸡WG、FI、CW、HEW、EW、BMW、BMY、LMW、AFW、AFY、 IFW、SFT;降低了F/G,对LMY有降低趋势。日粮添加100mg/kg GABA增加了22-42d热应激肉鸡WG、FI、EW、BMW、BMY、AFW、AFY、SFT;对CW、HEW、IFW有增加趋势。两种氨基酸在影响循环热应激肉鸡WG、FI、F/G、EW、EY、BMW、 BMY、AFW、AFY、IFW和SFT方面存在协同效应。在影响生产性能方面,Gln的作用具有较好的持续性,GABA在第1周时作用较强,随后明显减弱。
1.2循环热应激降低了28d、35d、42d肉鸡血清TP、葡萄糖、甘油三酯、Gin、Glu、T4、Ins水平、ALP活性,22d、35d血清GABA水平；增加了22d、35d、42d肉鸡血清CS、GN水平、GOT、GPT、LDH、CK、NOS活性。日粮添加5g/kg Gln增加了22d、35d、42d热应激肉鸡血清TP、Gln、Ins水平,降低了血清甘油三酯、CS、GN水平、GOT、CK、NOS活性。日粮添加100mg/kg GABA增加了22d、35d、42d热应激肉鸡血清TP、Gln、GABA、T3水平、ALP活性,对甘油三酯、Gln、T4、 CS、Ins、GN水平、GOT、GPT、LDH、CK、NOS活性的影响存在明显的时间特异性。两种氨基酸在影响循环热应激肉鸡血清TP、甘油三酯、GABA、Gln、T3、T4、 Ins、GN水平、ALP、LDH和CK活性方面存在明显的协同效应,在影响转氨酶(GOT、 GPT)和NOS方面仅在第1周时较显著。
1.3循环热应激增加了28、35、42d肉鸡胸肌pH、L*、水分、CP、CF、CA、 TL、F、MFDS,腿肌pH、L*、CA, DL、SF、MFDS;降低了肉鸡胸肌DL、CLMFDM,腿肌a*、b*、CP、CF、CL、TL、CL、MFDM。日粮添加5g/kg Gln降低了胸肌L*、SF、MFDS,腿肌L*、DL、SF、MFDS,增加了胸肌CP、CF、CA、TL、 DL、MFDM,腿肌a*、CP、CF、水分、CL、MFDM.日粮添加100mg/kg GABA降低了胸肌pH、TL、SF、腿肌pH、DL、SF,增加了胸肌a*、水分、CP、CF,腿肌a*、CF。两种氨基酸在影响循环热应激肉鸡胸肌pH、b*、水分、CP、WHC、TL、SF,腿肌pH、L*、a*、水分、CP、WHC、TL、SF方面存在明显的协同效应。此外,通过日粮Gln和GABA可以提高肌肉在冷藏过程中的色泽稳定性。
1.4循环热应激降低了28d、35d、42d肉鸡胸肌、腿肌的Gln、Glu、GABA含量、Glnase、GAD活性,增加了胸肌、腿肌的GS、GABA-T活性。日粮添加5g/kg Gln增加了胸肌的Gln、Glu含量、Glnase、GAD活性,腿肌的Gln、Glu、GABA含量、GAD活性；降低了胸肌、腿肌的GS活性,对胸肌的GABA含量、GABA-T活性、腿肌的Glnase、GABA-T活性有不同程度的影响。日粮添加100mg/kg GABA增加了胸肌的Gln、Glu含量、Glnase、GAD活性,腿肌Gln含量、GAD活性；降低了胸肌的GABA-T活性、腿肌的GS活性。两种氨基酸对循环热应激肉鸡胸肌、腿肌的Gln、 GABA含量、Glnase、GS、GABA-T活性的影响存在明显的协同效应。42d肉鸡胸肌、腿肌的Gln、GABA含量、Glnase、GS活性与L*、CP、CF、CL、MFDS和MFDM间存在显著的相关性。
1.5循环热应激降低了28d、35d、42d肉鸡胸肌、腿肌糖原、葡萄糖、乳酸含量,增加了胸肌、腿肌的AMP/ATP、AMPK含量、AMPK活性、HSP70含量、AMPKa2mRNA、HSP70mRNA表达量。日粮添加5g/kg Gln增加了胸肌、腿肌的HSP70含量、HSP70mRNA表达量、胸肌的葡萄糖、腿肌的糖原；降低了胸肌、腿肌的AMP/ATP、 AMPK含量、AMPK活性。日粮添加100mg/kg GABA增加胸肌的糖原、葡萄糖、腿肌的葡萄糖、乳酸,降低了胸肌、腿肌的AMP/ATP、AMPK活性、腿肌的HSP70含量、HSP70mRNA表达量。两种氨基酸对循环热应激肉鸡胸肌的葡萄糖、乳酸、腿肌的糖原、乳酸、胸肌和腿肌的AMP/ATP、AMPK活性、HSP70mRNA表达量存在明显的协同效应。42d肉鸡胸肌和腿肌的糖原、葡萄糖、乳酸与水分、CP、CF, AMP/ATP. AMPK含量、AMPK活性与L*、水分、CP、CF、WHC、DL、CL、MFDS和MFDM间,HSP70含量与WHC、TL、DL间存在较为显著的相关性。
2.体外试验：研究热应激影响家禽骨骼肌细胞HSP70、AMPK的调控关系及外源性Gln. GABA的干预作用。以1-2×106个/mL密度的成肌细胞进行试验。处理后置于37℃水浴中进行3h修复,收集细胞供分析。
2.1热应激条件下HSP70表达与AMPK活性变化时差性分析
共5组,每组6重复,基础培养基培养3h后进行热应激处理(42℃和45℃,0-120min)。结果表明：热应激提高了细胞的HSP70表达量和AMPK活性,以45℃处理的提高速度更快,程度更大,且以AMPK活性的变化速度快于HSP70表达量的变化。
2.2HSP70表达与AMPK活性问的调控关系分析
(1)QD抑制HSP70表达试验：共7组,每组6重复。用添加浓度为0-50umol/L的QD的基础培养基预培养3h,进行42℃和45℃、30min的热应激处理。结果表明：QD对HSP70表达的抑制作用存在明显的温度特异性,仅45℃时表现显著。同时,45℃时的AMPK活性具有不同程度的增加,显示HSP70能在一定程度上抑制/AMPK活性。
(2)AICAR激活AMPK试验：共7组,每组6重复。用添加浓度为0-0.5mmol/L的AICAR的基础培养基培养3h处理。结果表明：AICAR能有效激活适温条件下的骨骼肌细胞AMPK活性,但对HSP70的表达量无影响。
2.3Gln、GABA对细胞HSP70与AMPK的影响
各取5组,每组6重复。在基础培养基中分别添加0-20mmol/L Gln和0-0.8mmol/L GABA预培3h后,进行45℃,30mmin热应激处理。结果表明：Gln能有效增加热应激条件下骨骼肌细胞的HSP70表达量,同时抑制AMPK活性的增加。GABA能在一定程度上抑制AMPK活性的增加,但对HSP70表达量无影响。
2.4Gln、GABA对极端热应激骨骼肌细胞成活率的影响
各取3组,每组6重复。分别用添加0、10、20mmol/L Gln和0、0.4、0.8mmol/L GABA的基础培养基预孵3h后,采用48℃高温处理3h、6h,分析细胞成活率。结果表明：添加10、20mmol/L Gln提高了热应激处理3h、6h后的骨骼肌细胞成活率；添加0.4、0.8mmol/L GABA对细胞成活率无明显影响。
根据研究结果可以得出结论：
(1)本研究采用的30-34℃循环热应激对22-42d肉鸡生长性能、胴体性状、血清主要代谢物、激素和代谢酶等生化指标产生了显著的负面影响,也显示了机体对热应激的适应性。日粮添加5g/kg Gin、100mg/kg GABA能有效缓解循环热应激对上述指标带来的负面作用,同时表现出了明显的协同效应和时间特异性。
(2)从物质代谢角度分析,循环热应激显著影响了血清和骨骼肌中的Gln、Glu、 GABA水平,以及三种氨基酸间的代谢酶。通过日粮添加5g/kg Gln、100mg/kg GABA可以有效改善循环热应激条件下肉鸡骨骼肌中Gln、GABA的代谢状况,同时表现出了一定的协同效应和骨骼肌组织差异性。
(3)循环热应激明显造成了肉鸡机体的低能量状态,激活了骨骼肌中AMPK信号通路,促进了骨骼肌中HSP70转录和表达。通过日粮添加5g/kg Gln、100mg/kg GABA可以改善机体的能量状况,抑制骨骼肌AMPK信号通路,且表现出了一定的协同效应。同时说明,Gin的这种积极作用可能在一定程度上与促进HSP70表达有关,而GABA的改善作用与HSP70无关,可能与其神经系统和生理调控方面功能有关。
(4)循环热应激对胸肌、腿肌的化学组成和肉品质指标产生了显著的负面影响。通过日粮中添加5g/kg Gln、100mg/kg GABA对于缓解热应激对这些指标带来的负面影响具有积极作用,其中的机制与骨骼肌中Gin代谢变化、(?)AMPK信号通路激活状况、HSP70表达量的变化有密切关系,且表现出了一定的协同效应和骨骼肌组织差异性。
(5)通过体外骨骼肌细胞试验证明,热应激能激活肌细胞AMPK信号通路,促进HSP70的表达。热应激细胞中HSP70的表达可在一定程度上抑制AMPK信号通路的激活,起到保护细胞作用。在适温条件下细胞中AMPK的激活并不能导致HSP70表达量的变化。同时证明,外源性Gln、GABA均能抑制热应激骨骼肌细胞AMPK信号通路,但仅有Gln能有效保护热应激细胞,其作用与促进HSP70表达密切相关。
This experiment in vivo was conducted on broilers to evaluate the effects of circular heat stress (CHS) on growth performance, slaughter performance, carcass characteristic, serum biochemical parameter, meat quality, skeletal metabolism of Glutamine (Gln) and gamma-aminobutyric acid (GABA), heat shock protein (HSP70) and relative indexes in AMP-activated protein kinase (AMPK) signal path. The effects and interactions of dietary Gln and GABA on the broilers under CHS and the possible mechanism were also be evaluated. Moreover, the correlations between some skeletal indexes and meat quality indexes were analyzed. Another experiment in vitro was also conducted to evaluate regular relationship between HSP70and AMPK in poultry skeletal cell under heat stress (HS) and intervention of exogenous Gln and GABA on these indexes. Resolution of these issues will provide theoretical foundation for applying Gln and GABA against HS.
1. Experiment in vivo
The experiment was a2×2plus positive group (PC) of the treatments with two levels each of Gln (0and5g/kg Gln of diet) and GABA (0and100mg/kg GABA of diet). A total of36010-day-old healthy male Arbor Acres broiler chicks were procured and were housed from day10to day21in thermostatically controlled cages and were fed with same basic compound daily diet. On day22, after overnight (12h) feed withdrawal, chicks were wing-banded and randomly assigned to five treatment groups (6replicates of12birds per cage). The chicks of the four treatment groups (2×2) were housed in a circular high temperature controlled room (from9:00to14:00at30-34℃, then from14:00to18:00at34-30℃, and rest of the time at average23℃) while the PC group was housed in another optimal temperature controlled room (average23℃). Growth performance for each group was recorded from22to42days of age. At28.35,42days of age, three broilers from each cage with a body weight (BW) close to the group average was selected to analysis slaughter performance, carcass characteristics, serum biochemical parameters, meat quality, skeletal metabolism of Gin and GABA, HSP70and relative indexes in AMPK signal path, and to evaluate the correlations between some skeletal indexes and meat quality indexes. Results as below:
1.1CHS decreased weight gain (WG), feed intake (FI) of broiler from28d to42d old, carcass weight (CW), half-eviscerated weight (HEW), eviscerated weight (EW), breast muscle weight (BMW), breast muscle yield (BMY), leg muscle weight (LMW), abdominal fat weight (AFW), abdominal fat yield (AFY), intermuscle fat width (IFW)&subcutaneous fat thick (SFT) of42d old broiler, while increased feed-to-gain ratio (F/G)&leg muscle yield (LMY). The diet supplemented with5g/kg Gln increased WG, FI, CW, HEW, EW, BMW, BMY, LMW, AFW, AFY, IFW&SFT, while decreased F/G and had decreasing tendency for LMY of28-42d old broilers under CHS. The addition of100mg/kg GABA increased WG, FI, EW, BMW, BMY, AFW, AFY&SFT, while had increasing tendency for CW, HEW&IFW of28-42d old broilers under CHS. Different extent synergistic effects were found between Gln and GABA for WG, FI, F/G, EW, eviscerated yield (EY), BMW, BMY, AFW, AFY, IFW&SFT. With regard to the effects of Gln and GABA on growth performance,Gln had persistent action, while GABA had gradually weakened action.
1.2CHS decreased concentrations of TP, Glucose, triglyceride, Gln, Glu and T4, activities of ALP in serum of28,35,42d old broilers, and GABA content of28,35d old broilers, while increased concentrations of CS&GN, activities of GOT, GPT, LDH, CK&NOS in serum of28,35,42d old broilers. The diet supplemented with5g/kg Gln increased concentrations of TP, Gln&Ins, while decreased concentrations of triglyceride, CS&GN, activities of GOT, CK&NOS in serum of28,35,42d old broilers under CHS. The addition of100mg/kg GABA increased concentrations of TP, Gln, GABA&T3, ALP activities, while had obvious time-specifically effects on concentrations of triglyceride, Glu, T4, CS, Ins&GN, activities of GOT, GPT, LDH, CK&NOS in serum of28-42d old broilers under CHS. Different extent synergistic effects were found between Gln and GABA for serum concentrations of TP, triglyceride, GABA, Glu, T3, T4, Ins&GN, activities of ALP, LDH&CK, and GOT, GPT&NOS were found only in1st week under HS.
1.3CHS increased pH, L*, moisture, CP, CF, CA, TL, SF&muscle fiber density (MFDS) of breast muscle, pH, L*, CA, DL, SF&MFDS of leg muscle, while decreased DL, CL&muscle fiber diameter (MFDM) of breast muscle, a*, b*, CP, CF, CL, TL, CL&MFDM of leg muscle of28,35,42d old broilers. The diet supplemented with5g/kg Gln decreased L*, SF&MFDS of breast muscle, L*, DL, SF&MFDS of leg muscle, while increased CP, CF, CA, TL, DL&MFDM of breast muscle, a*, CP, CF, moisture CL&MFDM of leg muscle of broilers under CHS. The addition of100mg/kg GABA decreased pH, TL&SF breast muscle, pH, DL&SF of leg muscle, while increased a*, moisture, CP&CF of breast muscle, a*&CF of leg muscle of broilers under CHS. Different extent interactions were found between Gln and GABA for pH, b*, moisture, CP, WHC, TL&SF of breast muscle, pH, L*, a*, moisture, CP, WHC, TL&SF of leg muscle. Moreover, dietary Gln and GABA could improve color stability of breast and leg muscle during cold-storage.
1.4CHS decreased concentrations of Gln, Glu&GABA, activities of Glnase&GAD, while increased activities of GS&GABA-T in breast and leg muscle of28,35,42d old broilers. The diet supplemented with5g/kg Gin increased concentrations of Gln&Glu, activities of Glnase&GAD in breast muscle, concentrations of Gln, Glu&GABA, activities of GAD in leg muscle, while decreased GS activates of breast and leg muscle, and had different effects on GABA concentration and GABA-T activity in breast muscle, Glnase, GABA-T activities in leg muscle of broilers under CHS. The addition of100mg/kg GABA increased concentrations of Gln&Glu, activities of Glnase&GAD in breast muscle, Gln concentration and GAD activity in leg muscle, while decreased GABA-T activity in breast muscle and GS activity in leg muscle of broilers under CHS. Significant synergistic effects between dietary Gln and GABA on Gln content, GABA content, Glnase activity, GS activity&GABA-T activity were found. There were significant correlations among indexes of Gln content, GABA content, Glnase activity&GS activity and indexes of L*, CP, CF, CL, MFDS&MFDM in breast&leg muscle of42d broilers.
1.5CHS decreased concentrations of glycogen, Glucose&lactic acid, while increased AMP/ATP, AMPK content, AMPK activity, HSP70content, expression of AMPK a2mRNA and HSP70mRNA in breast and leg muscle of28,35,42d old broilers. The diet supplemented with5g/kg Gin increased HSP70content, HSP70mRNA expression in breast and leg muscle, glycogen content in breast muscle, Glucose content in leg muscle, while decreased AMP/ATP, AMPK content, AMPK activity in leg muscle of broilers under CHS. The addition of100mg/kg GABA increased glycogen and glucose contents in breast muscle, glucose and lactic acid contents in leg muscle, while decreased AMP/ATP and AMPK activity in breast and leg muscle, HSP70content, HSP70mRNA expression in leg muscle of broilers under CHS. Significant interactions between dietary Gln and GABA on glucose and lactic acid contents in breast muscle, glycogen and lactic acid in leg muscle, AMP/ATP, AMPK activity, HSP70mRNA expression in breast and leg muscle were found. There were significant correlations among indexes of glycogen, glucose&lactic acid and indexes of moisture, CP&CF, indexes of AMP/ATP, AMPK content&K activity and indexes of L*, moisture, CP, CF, WHC, DL, CL, MFDS&MFDM in breast&leg muscle. The significant correlations among HSP70and WHC, TL&DL were found.
2. Experiment in vitro
This experiment in vitro was conducted to evaluate the relationship between HSP70and AMPK in poultry skeletal cell under heat stress (HS), and the intervention of exogenous Gin and GAB A. The skeletal cell with density of1-2×106/ml were used in the experiments. After experimental treatment, the skeletal cell was repaired in37℃water bath for3h, and then was collected for analysis.
2.1Analysis on time difference of HSP70and AMPK under HS
A total of five treatment groups (6replicates of per group) were used in this experiment. HS (42℃and45℃,0-120min) was conducted on skeletal cell after culture in basic medium for3h. The results showed that HS increased HSP70expression and AMPK activity and the HS with45℃had more quickly and heavily increasing on the two indexes The increasing of AMPK activity was proceeding to HSP70expression.
2.2Analysis on regulation relationship between HSP70expression and AMPK activity
(1) Experiment about Quercetin (QD) inhibits HSP70expression:
A total of seven treatment groups (6replicates of per group) were used in this experiment. HS (42℃and45℃,30min) was conducted on skeletal cell after culture in basic medium with supplementation of0-50umol/L QD for3h. The results showed that inhibition of QD on HSP70expression showed obvious temperature-specificity, and the inhibition only displayed under45℃condition. Meanwhile, the increasing of AMPK activity under45℃showed that HSP70could inhibit AMPK activity to a certain extent.
(2) Experiment about AICAR activates AMPK:
A total of seven treatment groups (6replicates of per group) were used in this experiment. Skeletal cell was cultured in basic medium with supplementation of0-0.5mmol/L AICAT for3h under thermoneutrality. The results showed that AICAR could activate AMPK, but HSP70expression was not effected by AICAR.
2.3Effect of Gln and GABA on HSP70and AMPK
Two totals of five treatment groups (6replicates of per group) were used respectively in this experiment. HS (45℃,30min) was conducted on skeletal cell after cultured in basic medium supplemented with0-20mmol/L Gln or0-0.8mmol/L GABA for3h. The results showed that Gln could increase HSP70expression, while inhibit AMPK activity, and GABA could inhibit AMPK activity to a certain extent, but had no effect on HSP70expression in skeletal cell under HS.
2.4Effect of Gln and GABA on survival rate of skeletal cell under extremely HS
Two totals of three treatment groups (6replicates of per group) were used respectively in this experiment. HS (48℃,3h and6h) was conducted on skeletal cell after cultured in basic medium supplemented with0,10,20mmol/L Gln or0,0.4,0.8mmol/L GABA for3h. The results showed that10,20mmol/L Gln could increase survival rate of skeletal cell under HS for3h,6h (P<0.05or0.01), but0.4,0.8mmol/L GABA had no effect.
As stated above, the conclusions are as follows:
(1) The CHS (30-34℃) had obviously negative effects on growth performance, slaughter performance, carcass characteristics and serum biochemical parameters (main metabolites, hormones, metabolic enzymes), which also showed adaptability against HS. The diet supplemented with5g/kg Gln and100mg/kg GABA offered a potential nutritional strategy to prevent HS-related depression in these indexes of broiler, and also showed significant synergistic effect and time-specificity.
(2) From the material metabolism, CHS affected significantly Gln, Glu and GABA contents in serum and skeletal muscle and metabolic enzyme among them. Dietary Gln (5g/kg) and GABA (100mg/kg) improved metabolic state of Gln and GABA in skeletal muscle of broiler under CHS, and also showed a certain synergistic effect and skeletal tissue-variability.
(3) CHS obviously formed low-energy state of organism, activated AMPK signal path, and increased HSP70expression in skeletal muscle of broiler. Dietary Gln (5g/kg) and GABA (100mg/kg) improved energy state, inhibited AMPK signal path in skeletal muscle, and also showed a certain interactions between this two amino acids. The results also indicated that the positive action of Gln might be relevant to promoting HSP70expression. But the improvement of GABA was unrelated to HSP70expression, which may be involved in function of GABA in nervous system and physiological regulation.
(4) CHS had significant negative effects on chemical compositions and quality of breast and leg meat. Dietary Gln (5g/kg) and GABA (100mg/kg) offered a potential nutritional strategy to prevent HS-related depression in these indexes of broiler, and also showed a certain interaction and skeletal tissue-variability. The impossible mechanism of CHS affects meat quality is related to metabolic change of Gln, activated state of AMPK signal path, and variation of HSP70expression to agreater extent
(5) By test in vitro proved that HS activated AMPK signal path, and increased HSP70expression in skeletal muscle cell. Elevated HSP70expression could inhibit the activation of AMPK signal path in skeletal cell under HS and thus protect skeletal cell from HS-relative damage. The activation of AMPK could not affect HSP70expression in skeletal cell under thermoneutrality. Exogenous Gln and GABA could both inhibit activation of AMPK, but only Gln had effective protection to skeletal cell under HS, which was related to increasing HSP70expression.
引文
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