摘 要:殼寡糖是一種具有益生元活性的功能性寡糖,主要來(lái)源于蝦、蟹的外殼。作為抗生素潛在的替代品之一,殼寡糖在家禽生產(chǎn)中的開(kāi)發(fā)與應(yīng)用上有著巨大的潛力。本文在國(guó)內(nèi)外相關(guān)研究文獻(xiàn)的基礎(chǔ)上,重點(diǎn)闡述了殼寡糖的抗氧化、抗炎、調(diào)節(jié)腸道菌群、脂質(zhì)代謝及保護(hù)肝臟健康等方面的生物學(xué)功能及其在家禽養(yǎng)殖中的應(yīng)用,以期為殼寡糖在家禽生產(chǎn)中的研究和合理利用提供科學(xué)依據(jù)。
關(guān)鍵詞:殼寡糖;生物學(xué)功能;家禽
中圖分類號(hào):S816.7 文獻(xiàn)標(biāo)識(shí)碼:A 文章編號(hào):1673-1085(2024)10-0010-10
殼寡糖(Chitosan oligosaccharide,COS)是從蝦、貝類等甲殼類生物外殼中提取出來(lái)的物質(zhì)[1]。我國(guó)沿海地區(qū)的水產(chǎn)資源豐富,原本只能用作廢棄物的水生生物外殼,如果能加以合理利用,有望可以同時(shí)解決我國(guó)沿海地區(qū)資源浪費(fèi)和環(huán)境污染的雙重問(wèn)題。飼用抗生素能夠促進(jìn)家禽生長(zhǎng),控制一些疾病的產(chǎn)生和蔓延,但抗生素使用量過(guò)多,會(huì)導(dǎo)致產(chǎn)品中抗生素殘留及細(xì)菌耐藥性等諸多問(wèn)題[2]。研究和開(kāi)發(fā)新型替抗產(chǎn)品已成為飼料行業(yè)的發(fā)展趨勢(shì)。作為一種無(wú)污染、無(wú)殘留的天然新型飼料添加劑,COS具有抗氧化、抗炎、調(diào)節(jié)腸道菌群、脂質(zhì)代謝以及保護(hù)肝臟健康等多種生物學(xué)功能,具有替代抗生素的潛力。本文綜述了殼寡糖的生物學(xué)功能及其在家禽養(yǎng)殖中的應(yīng)用效果,以期為殼寡糖在家禽生產(chǎn)中的高效應(yīng)用提供科學(xué)依據(jù)。
1 殼寡糖的結(jié)構(gòu)
甲殼素是除纖維素外最豐富的天然資源,是蝦、蟹等甲殼類動(dòng)物外殼的主要成分[3]。甲殼素分子中由于存在大量的氫鍵而具有穩(wěn)定的結(jié)構(gòu),且乙?;潭瘸^(guò)90%,因此其溶解性差、難以被吸收利用[4]。殼聚糖是甲殼素脫乙酰度超過(guò)55%的產(chǎn)物,具有比甲殼素更好的生物相容性和溶解性,但仍不溶于水等一般的溶劑[5]。COS是由殼聚糖通過(guò)物理水解、酸水解或酶降解等方法解聚而成的[6]。低聚合度的COS含有活性官能團(tuán),即氨基和羥基,具有低分子量和粘度、較高的脫乙酰度和水溶性[7],這使其具有更顯著的生物學(xué)特性,如抗氧化、抗炎、調(diào)節(jié)腸道菌群、脂質(zhì)代謝以及保護(hù)肝臟健康等。COS的結(jié)構(gòu)式如圖 1所示。
2 殼寡糖的生物學(xué)功能
2.1 提高抗氧化能力
超氧化物歧化酶(Superoxide Dismutase,SOD)、谷胱甘肽過(guò)氧化物酶(Glutathione peroxidase,GSH-Px)的活性和丙二醛(Malonaldehyde,MDA)的含量是衡量動(dòng)物機(jī)體抗氧化系統(tǒng)強(qiáng)弱的重要指標(biāo),SOD、GSH-Px等抗氧化酶活性的升高和MDA含量的降低均能表明機(jī)體抗氧化能力有所提高[8]。Lan等[9]報(bào)道,在熱應(yīng)激條件下,飼糧中添加200 mg/kg COS能夠提高肉雞肝臟中SOD、GSH-Px等抗氧化酶的活性,降低MDA含量,表明COS可緩解熱應(yīng)激造成的氧化損傷。Xu等[10]報(bào)道,飼糧中添加75 g/kg COS能夠提高蛋雞血清總抗氧化能力,降低MDA含量。Lan等[11]報(bào)道,飼糧中添加600 mg/kg COS可增加麒麟雞肌肉中GSH-Px、SOD和過(guò)氧化氫酶(Catalase,CAT)活性,同時(shí)降低MDA含量,這表明COS能夠通過(guò)提高麒麟雞體內(nèi)抗氧化酶的活性來(lái)減輕氧化應(yīng)激引起的氧化損傷。Li等[12]報(bào)道,飼糧中添加COS可增加肉雞十二指腸SOD活性,降低空腸MDA含量,這表明COS能夠提高肉雞小腸的總抗氧化能力。也有研究發(fā)現(xiàn),COS的抗氧化功能可能與核因子E2相關(guān)因子2(Nuclear factor-erythroid 2-related factor 2,Nrf2)的活化有關(guān),即COS可通過(guò)促進(jìn)Nrf2的mRNA表達(dá)來(lái)增強(qiáng)抗氧化能力[13]。Chang等[14]報(bào)道,COS可以通過(guò)激活Nrf2/HO-1通路來(lái)提高GSH-Px的活性,從而緩解熱應(yīng)激誘導(dǎo)的黃羽肉雞胸肌的氧化損傷。飼糧中添加15 mg/kg COS可以提高脂多糖(Lipopolysaccharide,LPS)攻毒蛋雞腸道中Nrf2和HO-1的表達(dá),表明COS對(duì)LPS造成的氧化損傷具有保護(hù)作用[15]。由此可知,COS可能通過(guò)激活Nrf2通路來(lái)提高機(jī)體多種抗氧化酶活性及降低MDA含量,從而實(shí)現(xiàn)其抗氧化功能。
2.2 提高抗炎能力
核轉(zhuǎn)錄因子-κB(Nuclear factor-kappa B,NF-κB)是調(diào)節(jié)炎癥反應(yīng)的關(guān)鍵轉(zhuǎn)錄因子,可調(diào)控許多炎癥細(xì)胞因子的表達(dá)[16]。Yu等[17]報(bào)道,TLR4/MyD88/NF-κB信號(hào)通路被激活后,會(huì)促進(jìn)炎癥細(xì)胞因子的產(chǎn)生,因此可以通過(guò)抑制TLR4/NF-κB信號(hào)通路的激活來(lái)緩解炎癥反應(yīng)。在飼糧中添加COS可降低肉雞腸道中TLR4、MyD88、NF-κB p65、IL-1β的mRNA表達(dá)水平,提示COS可抑制TLR4/NF-κB信號(hào)通路的激活[18]。Lan等[18]報(bào)道,飼糧中添加COS有線性提高肉雞血清和脾臟中抗炎性細(xì)胞因子的趨勢(shì)。Deng等[19]報(bào)道,飼糧中添加COS能夠提高肉雞血清中抗炎性細(xì)胞因子IL-10的水平。此外,飼糧中添加COS還可顯著提高肉雞十二指腸中IL-10的水平,表明COS可從多種途徑提高肉雞的抗炎能力[20]。COS還可降低H2O2刺激的RAW264.7細(xì)胞中IL-1β、IL-6和TNF-α的mRNA表達(dá),表明COS具有緩解炎癥的作用[21]。Fathi等[22]報(bào)道,飼糧中添加COS顯著提高抗炎性細(xì)胞因子IL-10的水平,降低肉雞血清和肝臟中促炎性細(xì)胞因子TNF-α、IL-1β的水平。有研究發(fā)現(xiàn),LPS增加肉雞空腸IFN-γ及回腸IL-1β的水平,而COS可降低相應(yīng)腸段IFN-γ和IL-1β的水平,從而減輕LPS刺激對(duì)肉雞腸道的損害[15]。飼糧中添加COS能夠降低蛋雛雞血清中促炎性細(xì)胞因子IFN-γ、TNF-α的水平,說(shuō)明COS能改善家禽的抗炎能力[23]。以上結(jié)果表明,COS可通過(guò)降低促炎性因子水平和提高抗炎性因子水平兩種途徑,最終達(dá)到提高抗炎的目的。
2.3 調(diào)節(jié)腸道菌群
腸道菌群是由約1 014 種微生物組成的微生物區(qū)系,在維持腸道健康方面起著重要作用[24]。COS可作為腸道微生物的發(fā)酵底物來(lái)改善腸道微生物的組成,發(fā)酵產(chǎn)生的代謝產(chǎn)物又可調(diào)控腸道屏障功能以調(diào)節(jié)腸道微生物組成,從而更好地維持腸道系統(tǒng)的平衡[25]。徐晨希等[26]報(bào)道,飼糧中添加COS可提高肉雞盲腸中有益菌乳酸桿菌屬的相對(duì)豐度,降低致病菌糞腸桿菌屬的相對(duì)豐度。Zhang等[27]報(bào)道,飼糧中添加COS可提高肉雞盲腸中約翰遜乳桿菌、擬桿菌和沙拉亞硝基擬桿菌的含量,降低致病菌的相對(duì)豐度,表明COS能夠優(yōu)化腸道菌群的組成。Chang等[28]報(bào)道,飼糧中添加COS可顯著增加肉雞厚壁菌門和乳酸菌的豐度,降低變形菌的豐度,表明COS可以調(diào)節(jié)肉雞的腸道菌群。Menconi等[29]報(bào)道,鼠傷寒沙門氏菌可破壞肉雞腸道菌群平衡,降低有益菌的豐度,而COS改善腸道菌群平衡,降低鼠傷寒沙門氏菌的數(shù)量。飼糧中添加COS可在門水平上提高白羽王鴿回腸中變形菌門、放線菌門的豐度,在屬水平上提高假單胞菌屬和腸球菌屬等有益菌的豐度,降低羅斯氏菌屬等有害菌的豐度[30]。陳虹等[31]報(bào)道,飼糧中添加COS顯著提高鵪鶉盲腸內(nèi)容物中雙歧桿菌的數(shù)量,同時(shí)降低大腸桿菌的數(shù)量。由此可知,COS可通過(guò)提高有益菌的豐度及降低致病菌的豐度來(lái)改善腸道菌群,從而實(shí)現(xiàn)促進(jìn)腸道健康的功能。
2.4 調(diào)節(jié)脂質(zhì)代謝
研究表明,COS調(diào)節(jié)脂肪代謝的機(jī)制可能是通過(guò)調(diào)控與脂肪代謝相關(guān)基因的表達(dá)來(lái)影響脂肪代謝相關(guān)酶的活性,從而達(dá)到調(diào)節(jié)機(jī)體脂質(zhì)代謝的目的[32]。Xie等[33]報(bào)道,飼糧中添加COS可降低肝臟脂質(zhì)代謝基因的相對(duì)表達(dá)水平。盛東風(fēng)等[34]報(bào)道,飼糧中添加COS可下調(diào)乙酰輔酶A羧化酶(Acetyl coa carboxylase,ACC)和脂肪酸合成酶(Fatty acid synthase,F(xiàn)AS)的mRNA表達(dá)水平,顯著降低揚(yáng)州鵝肝臟中ACC的活性,說(shuō)明COS可通過(guò)調(diào)節(jié)與脂肪代謝相關(guān)基因和酶的活性來(lái)緩解肉雞腹部脂肪沉積。Wang等[35]報(bào)道,飼糧中添加COS顯著提高黃羽肉雞肝臟中脂蛋白脂肪酶(Lipoprotein lipase,LPL)的活性,表明COS能夠增強(qiáng)脂肪的分解代謝,有效減少肝臟脂肪沉積。Li等[36]報(bào)道,飼糧中添加COS顯著降低肉雞肝臟中FAS活性,提高肝臟中LPL活性,從而導(dǎo)致肉雞脂肪沉積減少。另外,COS還具有降血脂的作用。Wang等[35]報(bào)道,飼糧中添加COS會(huì)降低黃羽肉雞血清中甘油三酯(Triglycerides,TG)的含量,表明COS對(duì)體脂沉積有抑制作用。Li等[37]報(bào)道,飼糧中添加COS可提高肉雞血清高密度脂蛋白膽固醇(High density lipoprotein cholesterol,HDL-C)水平、降低TG和低密度脂蛋白膽固醇(Low-density lipoprotein cholesterol,LDL-C)水平。由此可知,殼寡糖調(diào)節(jié)脂質(zhì)代謝的方式不盡相同,具體作用途徑還有待進(jìn)一步研究。
2.5 保護(hù)肝臟健康
肝臟作為關(guān)鍵的代謝和解毒器官,對(duì)熱應(yīng)激敏感[38]。研究表明,熱應(yīng)激會(huì)引起氧化應(yīng)激和炎癥反應(yīng),從而誘發(fā)不同程度的肝損傷[39]。血液中谷草轉(zhuǎn)氨酶(Aspartate transaminase,AST)與谷丙轉(zhuǎn)氨酶(Alanine transaminase,ALT)的比值是反映動(dòng)物肝臟健康狀況的重要指標(biāo)。當(dāng)肝臟損傷時(shí),細(xì)胞膜通透性增加,肝臟中AST和ALT釋放到血液中,導(dǎo)致血液中AST和ALT活性增加[40]。慢性和急性熱應(yīng)激均可增加血清ALT和AST水平[41]。Lan等[42]報(bào)道,飼喂200 g/kg COS可顯著降低熱應(yīng)激肉雞血液中ALT、AST活性,說(shuō)明COS能夠減輕熱應(yīng)激對(duì)肉雞肝臟造成的損傷。Chang等[43]報(bào)道,熱應(yīng)激增加黃羽肉雞血清ALT和AST水平,表明熱應(yīng)激誘發(fā)了肝損傷。而飼糧中添加COS顯著降低熱應(yīng)激肉雞血清ALT和AST水平,表明COS對(duì)肉雞肝臟應(yīng)激損傷有明顯的保護(hù)作用。Lan等[44]報(bào)道,熱應(yīng)激可誘發(fā)肉雞肝臟的氧化應(yīng)激,增加血清中AST和ALT活性,飼糧中添加COS顯著降低血清AST和ALT活性,增加肝臟GSH-Px活性以及肝臟Nrf2、CAT、IL-10的表達(dá),說(shuō)明COS能夠減輕熱應(yīng)激對(duì)肉雞肝臟造成的氧化損傷,促進(jìn)肉雞的生長(zhǎng)。由此可知,COS能顯著降低動(dòng)物血清ALT、AST活力,改善肝功能,保護(hù)肝臟健康。
3 殼寡糖在家禽生產(chǎn)中的應(yīng)用
3.1 在雞生產(chǎn)中的應(yīng)用
平均日增重(Average daily weight gain,ADG)、平均日采食量(Average daily feed intake,ADFI)和飼料轉(zhuǎn)化率(Feed conversion ratio,F(xiàn)CR)是衡量機(jī)體生長(zhǎng)性能的重要指標(biāo)。大量研究報(bào)道,飼糧中添加不同水平的COS能夠改善雞的ADG、ADFI、FCR及養(yǎng)分消化率,見(jiàn)表1。Youssef等[45]報(bào)道,飼糧中添加100 mg/kg COS 顯著改善蛋雞產(chǎn)蛋率、蛋重和FCR。Osho等[46]報(bào)道,地塞米松(Dexamethasone,DEX)應(yīng)激可顯著增加肉雞體重?fù)p失,導(dǎo)致氧化應(yīng)激和免疫抑制,而飼糧中添加 100 mg/kg COS能夠緩解DEX對(duì)肉雞生長(zhǎng)性能的抑制作用,提高養(yǎng)分消化率。Lan等[47]報(bào)道,飼糧中添加 600 mg/kg COS 可改善卷曲雞的 ADG、ADFI和FCR,并提高肌肉中GSH-Px、SOD和CAT等抗氧化酶活性,同時(shí)降低MDA含量,上調(diào)Nrf2、HO-1等基因表達(dá)。Tao等[48]報(bào)道,在蛋雞脂肪肝綜合征模型中,飼糧中添加200、400、800 mg/kg COS可增強(qiáng)抗氧化酶活性,降低MDA水平,下調(diào)NF-κB、促炎細(xì)胞因子及促凋亡相關(guān)基因的mRNA表達(dá),進(jìn)而顯著提高產(chǎn)蛋率,降低料蛋比。Fathi等[49]報(bào)道,飼糧中添加300 mg/kg COS增加冷應(yīng)激肉雞血清和肝臟中IL-10水平,降低IL-1β和TNF-α水平,顯著改善肉雞ADG、FCR,有效緩解冷應(yīng)激導(dǎo)致的炎癥反應(yīng)。Lan等[18]報(bào)道,飼糧中添加200 mg/kg COS顯著提高熱應(yīng)激肉雞ADG、ADFI,增加血清中IL-10水平,同時(shí)降低IL-1β和TNF-α水平,緩解氧化應(yīng)激引起的炎癥反應(yīng)。Chang等[50]報(bào)道,飼糧中添加30 mg/kg COS顯著提高黃羽肉雞ADG、ADFI,增加回腸絨毛高度、絨毛高度與隱窩深度的比值,在一定程度上促進(jìn)營(yíng)養(yǎng)物質(zhì)的吸收。Xu等[51]報(bào)道,飼糧中添加75 mg/kg COS顯著提高豐達(dá)1號(hào)蛋雞產(chǎn)蛋量和FCR,顯著降低血清TG水平。Wang等[52]報(bào)道,飼糧中添加580 mg/kg COS顯著改善黃羽肉雞的FCR,提高LPL活性,促進(jìn)肝臟脂肪分解代謝,有效減少肝臟脂肪沉積。
3.2 在其他家禽生產(chǎn)中的應(yīng)用
COS除在雞以外的家禽中應(yīng)用較少,近年來(lái)關(guān)于國(guó)內(nèi)外COS在其他家禽生產(chǎn)上的研究結(jié)果見(jiàn)表2。陳中衛(wèi)等[62]報(bào)道,飼糧中添加40 mg/kg COS顯著改善櫻桃谷肉鴨ADG、ADFI、脾臟指數(shù)和胸腺指數(shù),顯著降低血清中D-乳酸水平,說(shuō)明COS能夠增強(qiáng)肉鴨的腸道屏障功能。Yuan等[63]報(bào)道,飼糧中添加1 200、2 400 mg/kg殼聚糖顯著改善肉鴨的ADG、ADFI和FCR,同時(shí)顯著提高免疫器官重量和淋巴細(xì)胞的增殖。宋濤等[64]報(bào)道,飼糧中添加200 g/t COS可改善北京肉鴨的ADG、ADFI和FCR,同時(shí)降低肌肉的滴水損失,而添加300 g/t COS明顯降低腹脂沉積,改善肉品質(zhì)。血清生化指標(biāo)是反映動(dòng)物生長(zhǎng)、機(jī)體代謝狀況的直接依據(jù),血清總蛋白和尿素氮含量反映了動(dòng)物體內(nèi)蛋白質(zhì)代謝和氨基酸的平衡情況[65]。辛清武等[66]報(bào)道,飼糧中添加40 mg/kg COS能提高北京肉鴨的ADG,同時(shí)提高血清中總蛋白含量,降低血清尿素、尿酸、膽固醇的含量,這表明COS能夠在一定程度上促進(jìn)北京鴨的生長(zhǎng)發(fā)育,提高其生產(chǎn)性能。另外,有報(bào)道表明,飼糧中添加40 mg/kg COS可顯著提高70日齡北京鴨血清SOD活性、GSH-Px活性,顯著降低血清中MDA含量,有效增強(qiáng)北京鴨的抗氧化能力[67]。Miao等[68]報(bào)道,日糧中添加200 mg/kg COS顯著提高豁眼鵝的免疫器官重量、血清免疫球蛋白、補(bǔ)體、激素和細(xì)胞因子水平,改善豁眼鵝的免疫功能??氯~艷等[69]報(bào)道,飼糧中添加3% COS可在一定程度上改善肉用鵪鶉的ADG和FCR,顯著降低其腹脂重量、脂肪表觀消化率和血清總膽固醇水平。綜上,飼糧中添加COS可以通過(guò)多種途徑直接或間接提高家禽的生長(zhǎng)性能,促進(jìn)家禽的生長(zhǎng)發(fā)育。
4 小結(jié)
當(dāng)前,飼用抗生素已被全面禁止,COS具有抗氧化、抗炎、調(diào)節(jié)腸道菌群和脂質(zhì)代謝等多種生物學(xué)功能,在家禽生產(chǎn)中具有廣泛的應(yīng)用前景。然而COS的結(jié)構(gòu)與其生物學(xué)功能之間的關(guān)系尚未完全被闡明,在未來(lái)還需要進(jìn)一步研究。此外,不同地區(qū)氣候環(huán)境、養(yǎng)殖品種、養(yǎng)殖模式存在較大差異,關(guān)于殼寡糖對(duì)不同養(yǎng)殖條件、不同生長(zhǎng)階段、不同家禽品種的作用效果以及添加劑量尚不清楚。因此,今后應(yīng)進(jìn)行深入研究。
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Biological Functions of Chitosan Oligosaccharide and Its Application in Poultry Production
Abstract: Chitosan oligosaccharide (COS) is a functional oligosaccharide with prebiotic activity, mainly derived from the shell of shrimp and crab. As one of the potential alternatives to antibiotics, chitosan oligosaccharide has great potential for development and application in poultry production. On the basis of relevant research literature, this paper focuses on the biological functions of chitosan oligosaccharide including antioxidant and anti-inflammatory activity, regulation of intestinal flora and lipid metabolism, protection of liver health and its application in poultry production, in order to provide a scientific basis for the research and rational utilization of chitosan oligosaccharide.
Keywords: Chitosan oligosaccharide; Biological functions; Poultry