摘要
日粮中的非纤维性碳水化合物（NFC）主要包括淀粉、可溶性糖以及中性洗涤可溶性纤维（NDSF），是反刍动物的主要能量来源。目前研究中，NFC常以单个值出现在日粮组成中，但不同来源饲料的NFC组成差异较大，例如玉米谷物中，NFC主要为淀粉；糖蜜中，NFC主要为可溶性糖（单糖和双糖）；而对于甜菜渣，NFC则主要为NDSF。然而，不同NFC组分在瘤胃中的发酵特性不同。与淀粉相比，果胶发酵不产生或产生极少乳酸，但具有较高的乙丙比，当瘤胃pH较低时，果胶发酵受到抑制或不再继续发酵；可溶性糖发酵较快，产生较高的丁酸。因而，在生产中将NFC作为统一整体考虑并不合适。生产中常用糖蜜和甜菜渣类饲料替代日粮中的部分谷物，但此类饲料对动物瘤胃发酵和生产性能的影响，各研究间结论并不一致。本研究通过四个人工瘤胃试验系统研究了体外环境条件下日粮NFC组分对养分降解、瘤胃发酵、微生物合成以及纤维分解菌菌群的影响，以期得到淀粉、可溶性糖和NDSF间适宜的供应方式，为糖蜜和甜菜渣类饲料在生产中的合理应用提供理论基础。现摘要如下：
试验一日粮瘤胃降解蛋白（RDP）水平和NFC类型对瘤胃发酵、微生物合成和纤维分解菌菌群的影响。本研究采用2×4的双因素试验设计，其中2种RDP水平，4种NFC类型（玉米淀粉、蔗糖、柑橘果胶和菊粉）。RDP水平通过向四种日粮中分别添加0（低RDP）或1.56g（高RDP）酪蛋白酸钠调节。主效应方面（2中RDP水平下的平均值，以下同），蔗糖和果胶组的干物质（DM）和有机物质（OM）表观消失率显著高于其它处理组（P <0.01）。日粮RDP水平和NFC来源在影响中性洗涤纤维（neutraldetergent fiber，NDF）的表观消失率上存在交互作用（P <0.01），低RDP处理组，蔗糖（P=0.10）和果胶组（P=0.09）趋向于低于淀粉组（P值），但高RDP处理组之间差异不显著。主效应方面，果胶组的液相和固相部分的Ruminococcus albus的16S rDNA拷贝数分别高于（P <0.01）和趋向于（P=0.10）高于淀粉组。在影响液相（P <0.01）和固相（P=0.02）Ruminococcus flavefaciens的16S rDNA拷贝数方面，日粮RDP水平和NFC来源间存在交互作用，其中低RDP下，蔗糖组固相Ruminococcus flavefaciens拷贝数趋向于（P=0.06）低于淀粉组。增加日粮RDP水平提高了所有处理组的总挥发性脂肪酸（VFA）产量（P <0.01）和总微生物氮（MN）流量（P <0.01）。主效应上，果胶组的乙酸摩尔比例（P <0.05）和乙丙比最高（P <0.01）。不论RDP水平，蔗糖和菊粉组的丁酸摩尔比例均高于其它处理组（P <0.01）。低RDP下各处理组的总MN流量差异不显著，但提高RDP使得蔗糖和果胶组的总MN流量分别高于（P <0.01）和趋向于（P=0.10）高于淀粉组。结果表明，日粮RDP水平和NFC类型在影响瘤胃发酵、微生物合成和纤维分解菌菌群方面存在交互作用，当日粮中有足够的瘤胃可利用氮时，蔗糖和果胶在微生物合成方面比淀粉更具有优势。
试验二日粮NDSF和蔗糖对瘤胃发酵、微生物合成和纤维分解菌菌群的影响。本研究采用2×2的双因素试验设计，日粮含有2种蔗糖添加量，高糖（约1.01g/d）和低糖（约0.26g/d）；2种NDSF水平，高NDSF（2.70g/d）和低NDSF（1.95g/d）。日粮NDSF和蔗糖在影响固相木聚糖酶活性（P=0.04）和DM （P=0.08）、OM （P=0.04）、NDF （P=0.01）、半纤维素（P=0.09）的表观消失率上存在交互作用，在高糖条件下增加NDSF供应量能够降低上述指标。尽管日粮NDSF和蔗糖在影响丁酸摩尔比例（P=0.05）方面存在交互作用，但与低NDSF日粮相比，高NDSF日粮增加了乙酸摩尔比例并降低了丁酸摩尔比例。然而，补加蔗糖对瘤胃VFA参数的影响则依赖日粮NDSF的水平。日粮蔗糖和NDSF在影响甲烷产量方面存在交互作用（P <0.01），提高NDSF供应量仅在低蔗糖日粮下增加甲烷产生量。提高NDSF和蔗糖供应量均降低了每天的净氨氮产量（P <0.01和P=0.03）。与低NDSF日粮相比，高NDSF日粮的总非氨态氮（NAN）（P=0.01）、MN（P=0.01）、固相微生物（SAM）（P <0.01）流量和微生物合成效率（EMS）（P=0.02）均较高。提高NDSF供应量增加了R. flavefaciens拷贝数（P <0.02），同时降低了R. albus拷贝数（P <0.01）。补加蔗糖提高了液相Ruminococcus albus拷贝数（P=0.01），但不影响总MN流量和固相纤维分解菌菌群。结果表明，在反刍动物日粮配合中，需要考虑NDSF和可溶性糖对瘤胃发酵以及纤维降解的交互影响；仍需进一步研究可溶性糖在瘤胃中的发酵和代谢。
试验三日粮NDSF替代淀粉对瘤胃发酵、微生物合成和纤维分解菌菌群的影响。本研究采用单因素试验设计，日粮的NDSF含量分别为12.7%、16.4%、20.1%和23.8%。提高日粮NDSF含量对DM（P=0.05）、OM（P=0.08）和NDF（P=0.06）表观消失率的影响，呈先增加后降低的趋势，较高值均出现在16.4%组。增加日粮NDSF含量提高了酸性洗涤纤维（ADF）（P=0.03）的表观消失率和纤维素酶活性，但降低了R. albus和R. flavefaciens的数量以及N的表观消失率（P <0.01）。用NDSF替代淀粉增加了乙酸摩尔比例（P <0.01）、乙丙比（P <0.01）、甲烷产量（P=0.03）和甲烷与VFA比（P<0.01）。提高日粮NDSF含量降低了氨氮产生量（P <0.01），增加了NAN（P <0.01）、SAM（P <0.01）流量和EMS （P=0.07），但总MN合成量在NDSF含量超过20.1%后没有进一步增加（P=0.07）。结果表明，当日粮NDSF含量占到16.4%时，提高了养分消化率（除N）和MN合成，进一步增加NDSF含量并不抑制MN合成，但显著降低了日粮N的消化。考虑到增加的微生物合成和瘤胃非降解N流量，用NDSF替代淀粉对反刍动物是有益的。
试验四高糖日粮NDSF替代淀粉对瘤胃发酵、微生物合成和纤维分解菌菌群的影响。本研究采用单因素试验设计，日粮的NDSF含量分别为12.9、16.6和20.3%。提高日粮NDSF含量降低了DM（P=0.03）、OM（P=0.05）、N（P <0.01）和NDF（P=0.09）的表观消失率，但增加了NFC的表观消失率（P=0.03）。用NDSF替代淀粉增加了丁酸摩尔比例（P <0.01）、乙丙比（P=0.06）以及甲烷与VFA比（P=0.04），降低了丙酸摩尔比例（P=0.04）。提高日粮NDSF含量降低了氨氮产量（P=0.04），增加了NAN（P=0.02）、SAM（P <0.01）流量和EMS（P=0.06），但总MN合成量在NDSF含量超过16.6%后没有进一步增加（P=0.13）。增加日粮NDSF含量降低了液相木聚糖酶和固相羧甲基纤维素酶活性，但不影响3种典型的纤维分解菌菌群。结果表明，高糖日粮下用NDSF替代淀粉虽降低了养分消化率（尤其日粮N），但提高了微生物合成。考虑到增加的微生物N合成和瘤胃非降解N流量，用NDSF替代淀粉对反刍动物是有益的，但需要考虑其对纤维消化的抑制影响。
Dietary non-fibre carbohydrates (NFC) are the major source of energy forhigh-producing dairy cattle. They are a very diverse group of carbohydrates, which includestarch, sugars, and neutral detergent soluble fiber (NDSF, largely pectic substances). Differentfeed sources vary greatly in the major components of NFC, although NFC has beenrepresented as a single value for feeds or diets. The NFC in corn grain is mostly starch, citruspulp provides sugars and NDSF, and sugars are predominant in molasses. However,fermentation of different NFC sources varies in digestion characteristics and the profiles oforganic acids produced. For example, pectins differ from starch and sugars producing little orno lactate, and they also elicit a higher acetate-to-propionate ratio compared with starch. Inaddition, the fermentation of pectin can be repressed or be ceased by lower ruminal pH.Sugars produce more butyrate compared with starch and pectin. Therefore, it is notappropriate for NFC as a whole. Effects of replacing cereal with molasses or sugar beet pulpon ruminal fermentation and animal production were not consistent among studies in pastyears. We investigate the effects of NFC on nutrients digestion, ruminal fermentation,microbial synthesis, and populations of ruminal cellulolytic bacteria using four artificialrumen (RUSITEC) experiments in order to determine the appropriate supply methods amongstarch, sugars, and NDSF and to provide theoretical basis for rational use of molasses andsugar beet pulp in the production and practice.
Trial1The objective of this study was to investigate the effects of rumen degradableprotein (RDP) and non-fibre carbohydrates (NFC) on ruminal fermentation, microbialsynthesis, and populations of ruminal cellulolytic bacteria using the rumen simulationtechnique (RUSITEC). Treatments consisted of four NFC types (corn starch, sucrose, pectin,and inulin) supplemented with0g/d (low RDP) or1.56g/d (high RDP) sodium caseinate.Apparent disappearance of dry matter (DM) and organic matter (OM) was greater (P <0.01)for the main effect means of sucrose and pectin than for other treatments, when averaged over both RDP supplementation rates. A NFC×RDP interaction (P <0.01) was observed forapparent neutral detergent fibre (NDF) disappearance, which tended to be lower for sucrose (P=0.10) and pectin (P=0.09) than for starch treatment under low RDP conditions, but did notdiffer among treatments under high RDP conditions. The16S ribosomal deoxyribonucleicacid (rDNA) copies of Ruminococcus albus were greater (P <0.01) and tended to be greater(P=0.10) for the main effect means of pectin than for starch treatment in liquid and solidfraction, respectively. There were NFC×RDP interactions for16S rDNA copy numbers ofRuminococcus flavefaciens, which in solid fraction tended to be lower (P=0.06) for sucrosethan for starch treatment under low RDP conditions. Increasing dietary RDP increased totalvolatile fatty acids production (P <0.01) and total microbial nitrogen (N) flow (P <0.01) inall treatments. The molar proportion of acetate and the ratio between acetate and propionatewere both greatest (P <0.05and P <0.01, respectively) for the main effect means of pectinamong treatments. Butyrate molar proportions were greater (P <0.01) for sucrose and inulintreatments than for other treatments regardless of RDP level. Total microbial N (MN) flow didnot differ among treatments under low RDP conditions, but sucrose (P <0.01) and pectin (P=0.10) produced greater MN than starch with increased RDP. Results showed NFC type, RDPlevel, and their interaction affected ruminal fermentation, microbial synthesis, and cellulolyticbacteria populations, and under sufficient ruminal available N sucrose and pectin had greateradvantage in microbial N synthesis than starch.
Trial2The objective of this study was to investigate the effects of neutral detergentsoluble fiber (NDSF) and sucrose supplementation on ruminal fermentation, microbialsynthesis, and populations of ruminal cellulolytic bacteria in Rusitec fermenters. Theexperiment had a2×2factorial design with two dosages of sucrose, low (ca.0.26g/d,low-sucrose) and high (ca.1.01g/d, high-sucrose), and two dosages of supplied NDSF, low(1.95g/d, low-NDSF) and high (2.70g/d, high-NDSF). Numerical or significant interactionsbetween NDSF and sucrose were detected for apparent disappearance of DM (P=0.08), OM(P=0.04), NDF (P=0.01), and hemicellulose (P=0.09), and xylanase from solid phase (P=0.04), which were reduced by supplemental NDSF only in high-sucrose diets. Althoughinteractions between NDSF and sucrose were detected for the molar proportions of butyrate(P=0.05), on the whole, high-NDSF diets increased the molar proportion of acetate andreduced that of butyrate compared with low-NDSF diets; however, the effects of supplementalsucrose on VFA profiles depended upon NDSF amount. A NDSF×sucrose interaction wasobserved for the production of methane (P <0.01) where supplementation with NDSF only increased the production of methane in low-sucrose diets. Supplementation with NDSF andsucrose both reduced the net production of ammonia-N (P <0.01and P=0.03, respectively).High-NDSF fermenters had greater daily flow of total non-ammonia N (NAN)(P=0.01),total MN (P=0.01), and solid-associated microbial pellets (SAM)(P <0.01) and greaterefficiency of microbial synthesis (EMS)(P=0.02), expressed as g microbial N/kg OMfermented compared with low-NDSF fermenters. Supplementation with NDSF resulted in anincrease in16S rDNA copies of Ruminococcus flavefaciens (P <0.02) and a reduction incopies of Ruminococcus albus (P <0.01). Supplementation with sucrose increased the16SrDNA copies of Ruminococcus albus (P=0.01) from liquid fraction, but did not affect dailytotal MN flow and cellulolytic bacterium populations from solid fraction. These data indicatethat the effects of the interaction between NDSF and sugars on ruminal fermentation and fiberdigestion should be taken into account in diet formulation. Ruminal fermentation andmetabolism of sugars warrant further investigation.
Trial3The objective of this study was to investigate the effects of replacing dietarystarch with neutral detergent soluble fiber (NDSF) on ruminal fermentation, microbialsynthesis, and populations of ruminal cellulolytic bacteria using the rumen simulationtechnique (RUSITEC). Experimental diets contained12.7,16.4,20.1, and23.8%NDSFsubstituted for dietary starch on a dry matter basis. With the increasing dietary NDSF,apparent disappearance of DM (P=0.05), OM (P=0.08) and NDF (P=0.06) had afluctuation change, increasing first and decreasing afterwards, with the maximum valueobserved in the16.4%NDSF diet. Increasing dietary NDSF caused an increase in apparentdisappearance of acid detergent fiber (ADF)(P=0.03) and fibrolytic enzyme activity and areduction in the number of ruminal cellulolytic bacteria (i.e., R. albus and R. flavefaciens) andapparent disappearance of dietary nitrogen (P <0.01). Substituting NDSF for starch appearedto increase the molar proportion of acetate (P <0.01), acetate: propionate (P <0.01), theproduction of methane (P=0.03), and methane: VFA (P <0.01). Increasing dietary NDSFreduced the daily production of ammonia-N (P <0.01) and increased the daily flow of NAN(P <0.01) and SAM (P <0.01), and EMS (P=0.07), but total microorganisms flow did notfurther increase as dietary NDSF exceeded20.1%of diet DM (P=0.07). Results suggest thatsubstituting NDSF for starch up to16.4%of diet DM increased digestion of nutrients (exceptfor N) and microbial synthesis, and further increase (from16.4to23.8%) in dietary NDSF didnot repress microbial synthesis but significantly reduced digestion of dietary N. Replacingdietary starch with NDSF may be beneficial for ruminants production due to increased flow of microbial protein and ruminally undegradable feed protein.
Trial4The objective of this study was to investigate the effects of replacing dietarystarch with neutral detergent soluble fiber (NDSF) in high sucrose diets on ruminalfermentation, microbial synthesis, and populations of ruminal cellulolytic bacteria using therumen simulation technique (RUSITEC). Experimental diets contained12.9,16.6, and20.3%NDSF substituted for dietary starch on a dry matter basis. Increasing dietary NDSF reducedor tended to reduce the apparent disappearance of DM (P=0.03), OM (P=0.05), N (P <0.01), and NDF (P=0.09), but increased that of NFC (P=0.03). Substituting NDSF forstarch appeared to increase the molar proportion of butyrate (P <0.01), acetate: propionate(P=0.06), and methane: VFA (P=0.04), and reduced the molar proportion of propionate (P=0.04). Increasing dietary NDSF reduced the daily production of ammonia-N (P=0.04) andincreased the daily flow of NAN (P=0.02) and SAM (P <0.01), and EMS (P=0.06). Totalmicroorganisms flow did not further increase as dietary NDSF exceeded16.6%of diet DM (P=0.13). Increasing dietary NDSF caused a reduction in xylanase activity from liquid fraction(P=0.02) and carboxymethylcellulase activity from solid fraction (P=0.02), but did notaffect typical cellulolytic bacteria population. Results suggest that substituting NDSF forstarch reduced digestion of nutrients especially dietary N, but improved the microbialsynthesis under high sugar conditon. Replacing dietary starch with NDSF may be beneficialfor ruminant production due to increased flow of microbial protein and ruminallyundegradable feed protein, but its repression on fiber digestion should be taken into account indiet formulation.
引文
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