Skip to main content Accessibility help

High-grain feeding alters caecal bacterial microbiota composition and fermentation and results in caecal mucosal injury in goats

  • Junhua Liu (a1), Tingting Xu (a1), Weiyun Zhu (a1) and Shengyong Mao (a1)


The effect of high-grain (HG) feeding on caecal bacterial microbiota composition and fermentation and mucosa health is largely unknown. In the present study, ten male goats were randomly assigned to either a group fed a hay diet (0 % grain; n 5) or a group fed a HG diet (65 % grain; n 5) to characterise the changes in the composition of the bacterial community and mucosal morphology in the caecum. After 7 weeks of feeding, the HG diet decreased the caecal pH (P< 0·001) and increased (P< 0·001 to P< 0·004) the caecal digesta concentrations of total volatile fatty acids and lipopolysaccharide (LPS). Pyrosequencing of the 16S ribosomal RNA gene revealed that HG feeding increased (P= 0·001 to P= 0·009) the abundance of predominant genera Turicibacter and Clostridium in the caecal lumen and in the caecal mucosa and decreased (P< 0·001 to P< 0·009) the proportion of Bacteroides in the lumen and Mucispirillum in the mucosa compared with the hay diet. Furthermore, the HG diet-fed goats exhibited intense epithelial damage and up-regulation (P< 0·001 to P< 0·025) of the relative mRNA expression of IL-1β, IL-6, IL-12 and interferon-γ (IFN-γ) in the caecal mucosa. The correlation analysis revealed that alterations in caecal pH, LPS concentration and mucosa-associated microbiota abundance during HG feeding might partly contribute to local inflammation. Collectively, these results provide insight into the adaptive response of caecal bacterial populations to HG feeding in goats and reveal that the fermentable substrates that flow into the caecum may cause dramatic alterations in microbial compositions and play a significant role in caecal dysfunction.

  • View HTML
    • Send article to Kindle

      To send this article to your Kindle, first ensure is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about sending to your Kindle. Find out more about sending to your Kindle.

      Note you can select to send to either the or variations. ‘’ emails are free but can only be sent to your device when it is connected to wi-fi. ‘’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

      Find out more about the Kindle Personal Document Service.

      High-grain feeding alters caecal bacterial microbiota composition and fermentation and results in caecal mucosal injury in goats
      Available formats

      Send article to Dropbox

      To send this article to your Dropbox account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Dropbox.

      High-grain feeding alters caecal bacterial microbiota composition and fermentation and results in caecal mucosal injury in goats
      Available formats

      Send article to Google Drive

      To send this article to your Google Drive account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Google Drive.

      High-grain feeding alters caecal bacterial microbiota composition and fermentation and results in caecal mucosal injury in goats
      Available formats


Corresponding author

* Corresponding author: S. Mao, fax +86 25 84395314, email


Hide All
1 Liu, JH, Xu, TT, Liu, YJ, et al. (2013) A high-grain diet causes massive disruption of ruminal epithelial tight junctions in goats. Am J Physiol Regul Integr Comp Physiol 305, R232R241.
2 Klevenhusen, F, Hollmann, M, Podstatzky-Lichtenstein, L, et al. (2013) Feeding barley grain-rich diets altered electrophysiological properties and permeability of the ruminal wall in a goat model. J Dairy Sci 96, 22932302.
3 Steele, MA, Croom, J, Kahler, M, et al. (2011) Bovine rumen epithelium undergoes rapid structural adaptations during grain-induced subacute ruminal acidosis. Am J Physiol Regul Integr Comp Physiol 300, R1515R1523.
4 Gressley, TF, Hall, MB & Armentano, LE (2011) Ruminant nutrition symposium: productivity, digestion, and health responses to hindgut acidosis in ruminants. J Anim Sci 89, 11201130.
5 Faichney, GJ (1968) Volatile fatty acids in the caecum of the sheep. Aust J Biol Sci 21, 177180.
6 Dixon, RM & Nolan, JV (1982) Studies of the large intestine of sheep. 1. Fermentation and absorption in sections of the large intestine. Br J Nutr 47, 289300.
7 Li, S, Khafipour, E, Krause, DO, et al. (2012) Effects of subacute ruminal acidosis challenges on fermentation and endotoxins in the rumen and hindgut of dairy cows. J Dairy Sci 95, 294303.
8 Metzler-Zebeli, BU, Schmitz-Esser, S, Klevenhusen, F, et al. (2013) Grain-rich diets differently alter ruminal and colonic abundance of microbial populations and lipopolysaccharide in goats. Anaerobe 20, 6573.
9 Steele, MA, AlZahal, O, Hook, SE, et al. (2009) Ruminal acidosis and the rapid onset of ruminal parakeratosis in a mature dairy cow: a case report. Acta Vet Scand 51, 39.
10 Emmanuel, DGV, Madsen, KL, Churchill, TA, et al. (2007) Acidosis and lipopolysaccharide from Escherichia coli B: 055 cause hyperpermeability of rumen and colon tissues. J Dairy Sci 90, 55525557.
11 Chin, AC, Flynn, AN, Fedwick, JP, et al. (2006) The role of caspase-3 in lipopolysaccharide-mediated disruption of intestinal epithelial tight junctions. Can J Physiol Pharmacol 84, 10431050.
12 Plaizier, JC, Khafipour, E, Li, S, et al. (2012) Subacute ruminal acidosis (SARA), endotoxins and health consequences. Anim Feed Sci Technol 172, 921.
13 Crawford, C, Sepulveda, MF, Elliott, J, et al. (2007) Dietary fructan carbohydrate increases amine production in the equine large intestine: implications for pasture-associated laminitis. J Anim Sci 85, 29492958.
14 Qin, WL (1982) Determination of rumen volatile fatty acids by means of gas chromatography. J Nanjing Agric Coll 4, 110116.
15 Barker, SB & Summerson, WH (1941) The colorimetric determination of lactic acid in biological material. J Biol Chem 138, 535554.
16 Mao, SY, Zhang, G & Zhu, WY (2008) Effect of disodium fumarate on ruminal metabolism and rumen bacterial communities as revealed by denaturing gradient gel electrophoresis analysis of 16S ribosomal DNA. Anim Feed Sci Technol 140, 293306.
17 Quince, C, Lanzen, A, Davenport, RJ, et al. (2011) Removing noise from pyrosequenced amplicons. BMC Bioinformatics 12, 38.
18 Pruesse, E, Quast, C, Knittel, K, et al. (2007) SILVA: a comprehensive online resource for quality checked and aligned ribosomal RNA sequence data compatible with ARB. Nucleic Acids Res 35, 71887196.
19 Wang, Q, Garrity, GM, Tiedje, JM, et al. (2007) Naive Bayesian classifier for rapid assignment of rRNA sequences into the new bacterial taxonomy. Appl Environ Microbiol 73, 52615267.
20 Lozupone, C & Knight, R (2005) UniFrac: a new phylogenetic method for comparing microbial communities. Appl Environ Microbiol 71, 82288235.
21 Snipes, RL (1981) Anatomy of the cecum of the laboratory mouse and rat. Anat Embryol 162, 455474.
22 Chomczynski, P & Sacchi, N (1987) Single-step method of RNA isolation by acid guanidinium thiocyanate–phenol–chloroform extraction. Anal Biochem 162, 156159.
23 Vorachek, WR, Bobe, G & Hall, JA (2013) Reference gene selection for quantitative PCR studies in bovine neutrophils. Adv Biosci Biotechnol 4, 614.
24 Clark, JH & Davis, CL (1980) Some aspects of feeding high producing dairy cows. J Dairy Sci 63, 873885.
25 Owens, FN, Secrist, DS, Hill, WJ, et al. (1998) Acidosis in cattle: a review. J Anim Sci 76, 275286.
26 Huntington, GB (1997) Starch utilization by ruminants: from basics to the bunk. J Anim Sci 75, 852867.
27 Owens, FN, Zinn, RA & Kim, YK (1986) Limits to starch digestion in the ruminant small intestine. J Anim Sci 63, 16341648.
28 Romero-Perez, GA, Ominski, KH, McAllister, TA, et al. (2011) Effect of environmental factors and influence of rumen and hindgut biogeography on bacterial communities in steers. Appl Environ Microbiol 77, 258268.
29 Godoy-Vitorino, F, Goldfarb, KC, Karaoz, U, et al. (2012) Comparative analyses of foregut and hindgut bacterial communities in hoatzins and cows. ISME J 6, 531541.
30 Lu, L & Walker, WA (2001) Pathologic and physiologic interactions of bacteria with the gastrointestinal epithelium. Am J Clin Nutr 73, 1124S1130S.
31 Malmuthuge, N, Griebel, PJ & Guan le, L (2014) Taxonomic identification of commensal bacteria associated with the mucosa and digesta throughout the gastrointestinal tracts of preweaned calves. Appl Environ Microbiol 80, 20212028.
32 Zhang, C, Zhang, M, Wang, S, et al. (2010) Interactions between gut microbiota, host genetics and diet relevant to development of metabolic syndromes in mice. ISME J 4, 232241.
33 Di Rienzi, SC, Sharon, I, Wrighton, KC, et al. (2013) The human gut and groundwater harbor non-photosynthetic bacteria belonging to a new candidate phylum sibling to Cyanobacteria. eLife 2, e01102.
34 Ley, RE, Backhed, F, Turnbaugh, P, et al. (2005) Obesity alters gut microbial ecology. Proc Natl Acad Sci U S A 102, 1107011075.
35 Delgado, S, Cabrera-Rubio, R, Mira, A, et al. (2013) Microbiological survey of the human gastric ecosystem using culturing and pyrosequencing methods. Microb Ecol 65, 763772.
36 Guinane, CM, Tadrous, A, Fouhy, F, et al. (2013) Microbial composition of human appendices from patients following appendectomy. MBio 4, e00366e00412.
37 Mao, SY, Zhang, RY, Wang, DS, et al. (2013) Impact of subacute ruminal acidosis (SARA) adaptation on rumen microbiota in dairy cattle using pyrosequencing. Anaerobe 24, 1219.
38 Shanks, OC, Kelty, CA, Archibeque, S, et al. (2011) Community structures of fecal bacteria in cattle from different animal feeding operations. Appl Environ Microbiol 77, 29923001.
39 Ahn, JH, Hong, IP, Bok, JI, et al. (2012) Pyrosequencing analysis of the bacterial communities in the guts of honey bees Apis cerana and Apis mellifera in Korea. J Microbiol 50, 735745.
40 Roesch, LF, Lorca, GL, Casella, G, et al. (2009) Culture-independent identification of gut bacteria correlated with the onset of diabetes in a rat model. ISME J 3, 536548.
41 Gagnon, N, Talbot, G, Ward, P, et al. (2007) Evaluation of bacterial diversity in the gut of piglets supplemented with probiotics using ribosomal intergenic spacer analysis. Can J Anim Sci 87, 207219.
42 Bosshard, PP, Zbinden, R & Altwegg, M (2002) Turicibacter sanguinis gen. nov., sp. nov., a novel anaerobic, Gram-positive bacterium. Int J Syst Evol Microbiol 52, 12631266.
43 Garcia, JP, Adams, V, Beingesser, J, et al. (2013) Epsilon toxin is essential for the virulence of Clostridium perfringens type D infection in sheep, goats, and mice. Infect Immun 81, 24052414.
44 Azcarate-Peril, MA, Foster, DM, Cadenas, MB, et al. (2011) Acute necrotizing enterocolitis of preterm piglets is characterized by dysbiosis of ileal mucosa-associated bacteria. Gut Microbes 2, 234243.
45 McLellan, SL, Newton, RJ, Vandewalle, JL, et al. (2013) Sewage reflects the distribution of human faecal Lachnospiraceae. Environ Microbiol 15, 22132227.
46 Tajima, K, Aminov, RI, Nagamine, T, et al. (2001) Diet-dependent shifts in the bacterial population of the rumen revealed with real-time PCR. Appl Environ Microbiol 67, 27662774.
47 Robertson, BR, O'Rourke, JL, Neilan, BA, et al. (2005) Mucispirillum schaedleri gen. nov., sp. nov., a spiral-shaped bacterium colonizing the mucus layer of the gastrointestinal tract of laboratory rodents. Int J Syst Evol Microbiol 55, 11991204.
48 Van den Abbeele, P, Belzer, C, Goossens, M, et al. (2013) Butyrate-producing Clostridium cluster XIVa species specifically colonize mucins in an in vitro gut model. ISME J 7, 949961.
49 Nagaraja, TG, Bartley, EE, Fina, LR, et al. (1978) Relationship of rumen Gram-negative bacteria and free endotoxin to lactic acidosis in cattle. J Anim Sci 47, 13291337.
50 Gozho, GN, Plaizier, JC, Krause, DO, et al. (2005) Subacute ruminal acidosis induces ruminal lipopolysaccharide endotoxin release and triggers an inflammatory response. J Dairy Sci 88, 13991403.
51 Bertok, L (1998) Effect of bile acids on endotoxin in vitro and in vivo (physico-chemical defense). Bile deficiency and endotoxin translocation. Ann N Y Acad Sci 851, 408410.
52 Ribeiro, MM, Xu, X, Klein, D, et al. (2010) Endotoxin deactivation by transient acidification. Cell Transplant 19, 10471054.
53 Mortensen, PB & Clausen, MR (1996) Short-chain fatty acids in the human colon: relation to gastrointestinal health and disease. Scand J Gastroenterol Suppl 216, 132148.
54 Andrews, F, Buchanan, B, Elliot, S, et al. (2005) Gastric ulcers in horses. J Anim Sci 83, E18E21.
55 Thibault, R, Blachier, F, Darcy-Vrillon, B, et al. (2010) Butyrate utilization by the colonic mucosa in inflammatory bowel diseases: a transport deficiency. Inflamm Bowel Dis 16, 684695.
56 Saleh, M & Trinchieri, G (2011) Innate immune mechanisms of colitis and colitis-associated colorectal cancer. Nat Rev Immunol 11, 920.
57 Elinav, E, Strowig, T, Kau, AL, et al. (2011) NLRP6 inflammasome regulates colonic microbial ecology and risk for colitis. Cell 145, 745757.
58 Kellermayer, R, Dowd, SE, Harris, RA, et al. (2011) Colonic mucosal DNA methylation, immune response, and microbiome patterns in Toll-like receptor 2-knockout mice. FASEB J 25, 14491460.
59 Lupp, C, Robertson, ML, Wickham, ME, et al. (2007) Host-mediated inflammation disrupts the intestinal microbiota and promotes the overgrowth of Enterobacteriaceae. Cell Host Microbe 2, 204.
60 Khafipour, E, Krause, DO & Plaizier, JC (2009) A grain-based subacute ruminal acidosis challenge causes translocation of lipopolysaccharide and triggers inflammation. J Dairy Sci 92, 10601070.
61 Wang, DS, Zhang, RY, Zhu, WY, et al. (2013) Effects of subacute ruminal acidosis challenges on fermentation and biogenic amines in the rumen of dairy cows. Livest Sci 155, 262272.
62 Plaizier, JC, Krause, DO, Gozho, GN, et al. (2008) Subacute ruminal acidosis in dairy cows: the physiological causes, incidence and consequences. Vet J 176, 2131.
63 Mainardi, SR, Hengst, BA, Nebzydoski, SJ, et al. (2011) Effects of abomasal oligofructose on blood and feces of Holstein steers. J Anim Sci 89, 25102517.


Type Description Title
Supplementary materials

Liu Supplementary Material
Tables S1-S5 and Figures S1-S6

 Word (6.5 MB)
6.5 MB


Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed