McSweeney CS, Palmer B, McNeill DM, Krause DO. Hrassnigg N, Crailsheim K. Differences in drone and worker physiology in honeybees (. These theoretical distinctions explain our separation of sections of this review devoted to digesters that rely largely on intrinsic enzymes to digest relatively nonrefractory materials in foods and sections devoted to digesters that typically ferment relatively refractory materials with the aid of symbiotic microbes. Heart. The three herbivores circled are individuals of red and giant panda, which are members of the order Carnivora. Identify structures that are a part of the digestive system, respiratory system, circulatory system, reproductive system, and excretory system. The digestive lysozyme is expressed in the acidic compartment of the foregut, has an acidic pH optimum, and is relatively resistant to breakdown by pepsin [reviewed by reference (303)]. Bars (i.e., means) within a discrete time period (i.e., at 6, 24, 48, or 72 h) that share a common letter did not differ significantly, whereas different letters indicate significant differences at P < 0.05. Ecologia Nutricional de Insetos e Suas Implicacoes no Manejo de Pragas. Coffee leaf miner trypsin inhibition with castor bean leaf extracts mediated by a non-protein agent. Stein ED, Diamond JM. Rumination has evolved independently in the ruminants and camels; kangaroos display more irregular cycles of regurgitation/swallowing that is known as merycism. 18B). The mechanistic basis of the impact of diet on digestive enzyme activity has not been investigated in most species but, where studied, there is persuasive evidence that differential enzyme activity is underpinned by changes in gene expression. Scharf ME, Kovaleva ES, Jadhao S, Campbell JH, Buchman GW, Boucias DG. Oxidative metabolism in the locust rectum. The usnic acid-resistant microbe is one of at least three fairly well-documented examples of ruminal microorganisms that can apparently tolerate some SMs. Harig JM, Ng EK, Dudeja PK, Brasitus TA, Ramaswamy K. Transport of n-butyrate into human colonic luminal membrane vesicles. Sklan D, Geyra A, Tako E, Gal-Gerber O, Uni Z. Ontogeny of brush border carbohydrate digestion and uptake in the chick. In addition, once the chyme leaves the stomach, the material is quite fluid in consistency. Bifano TD, Samuels RI, Alexandre D, Silva CP. There are four basic types of digestive systems: monogastric, avian, rumi- nant, and pseudo-ruminant. Chang MH, Chediack JG, Caviedes-Vidal E, Karasov WH. Metagenomic discovery of biomass-degrading genes and genomes from cow rumen. The areas under the curves (AUCs) are used to calculate fractional absorption, f, which averaged 87 3%. Increases in both SI activity and glucose transport occurred 2 days before hatch and at hatch day. Recent studies with fish, birds, and mammals exemplify these improvements. Tannin-binding proteins in saliva of deer and their absence in saliva of sheep and cattle. Studies using rat, mouse, and human fetal intestine grafted into adult hosts, or using altered diets, have shown that many of these changes occur in the absence of specific ontogenetic signals from either the lumen or circulation. Lysozyme [hydrolyzes peptidoglycan in G(+) bacterial cell walls (. Getachew G, Pittroff W, Putnam DH, Dandekar A, Goyal S, DePeters EJ. Also, work with pigs (438) and humans (168) that do not reingest feces demonstrates that there is another unknown pathway for absorption of microbially produced essential nutrients. The proton-coupled amino acid transporter, SLC36A1 (hPAT1), transports Gly-Gly, Gly-Sar and other Gly-Gly mimetics. The complexed tannins may escape both enzymatic and microbial degradation, and may be excreted in the feces, thus protecting the animal from either damage to the gut epithelium, true digestibility reduction, or toxicity (11). Chang MH, Karasov WH. Comparison of gastrointestinal transit times between chickens from D+ and D- genetic lines selected for divergent digestion efficiency. Srinivasan A, Giri AP, Gupta VS. Arts ICW, Sesink ALA, Hollman PCH. Buchon N, Broderick NA, Chakrabarti S, Lemaitre B. Invasive and indigenous microbiota impact intestinal stem cell activity through multiple pathways in Drosophila. The discovery of efflux transporters over the past 2 to 3 decades across many animal phyla revealed another process by which passive absorption of lipophillic SMs might be limited. Shu R, David ES, Ferraris RP. Two processes can mediate increased transporter function: recruitment of preexisting transporter protein in the cytoplasm to the membrane (as occurs for GLUT2 in response to dietary glucose, see Section Absorption of carbohydrates), and elevated gene expression. Another advantage of paracellular absorption is that it is an energetically cheap way to match absorption rate to substrate concentration in the diet and lumen. The esophagus,stomach,liver . These SMs are thus stored in an inactive form until activated by a glycohydrolase enzyme (e.g., -glucosidase). Enattah NS, Jensen TGK, Nielsen M, Lewinski R, Kuokkanen M, Rasinpera H, El-Shanti H, Seo JK, Alifrangis M, Khalil IF, Natah A, Ali A, Natah S, Comas D, Mehdi SQ, Groop L, Vestergaard EM, Imtiaz F, Rashed MS, Meyer B, Troelsen J, Peltonen L. Independent introduction of two lactase-persistence alleles into human populations reflects different history of adaptation to milk culture. Coll M, Guershon M. Omnivory in terrestrial arthropods: Mixing plant and prey diets. Zhang HZ, Malo C, Buddington RK. Lalles JP. Another famous example is the bacterium Synergistes jonesii, which is capable of degrading mimosine metabolites and imparts mimosine resistance in the host ruminant, allowing it to eat Leucaena spp. 12). These changes are predicted by the integrative model [Eq. An organ system is a network of individual organs that work with each other for a single purpose in the body. Austin PJ, Suchar LA, Robbins CT, Hagerman AE. Laino A, Cunningham ML, Garcia F, Heras H. First insight into the lipid uptake, storage and mobilization in arachnids: Role of midgut diverticula and lipoproteins. Expression of Na+/glucose co-transporter 1 (SGLT1) in the intestine of piglets weaned to different concentrations of dietary carbohydrate. 5 of reference (43).]. Meleshkevitch EA, Assis-Nascimento P, Popova LB, Miller MM, Kohn AB, Phung EN, Mandal A, Harvey WR, Boudko DY. Goel G, Puniya AK, Aguilar CN, Singh K. Interaction of gut microflora with tannins in feeds. Intestinal enzymes can activate certain toxins. Other mites that eat and grow on bacteria have higher activity levels of lysozyme, which breaks down bacterial cell walls (141). But, one of the congeneric but obligate carnivores (Xiphister atropurpureus) also increased -amylase without a diet shift, which suggested that phylogeny plays a role. Within each figure, points that share the same lower case letters do not differ significantly in mean value [Fig. We distinguish the term absorption (transport from gut lumen to body tissues by either the paracellular or transcellular route) from uptake, which refers to the transport from the gut lumen across the apical membrane of the gut epithelial cell (one step in transcellular transport). Physiological and Ecological Adaptations to Feeding In Vertebrates. Coexpression of ATP-binding cassette proteins ABCG5 and ABCG8 permits their transport to the apical surface. As a general rule, digestive efficiency on a food type declines with increasing amount of refractory material in food. The tips of the microvilli form web-type structures called glycocalyx.Amino acids and simple sugars released into the brush border membrane are absorbed into the microvilli first, then into the villi, and then pass into the circulatory system. Elvin-Lewis M. Should we be concerned about herbal remedies. Irritation in this area due to fine particle size, stress or other environmental factors can contribute to ulcer formation in swine. In humans and other mammals, all regions of the GI tract are colonized, including the highly acidic stomach, which bears a diverse community of bacteria and some fungi (30). Thus, key digestive adaptations of most herbivores besides special compartment(s) to maintain a microbiota are adjustments in digestive compartment sizes and possession of other GI structures that slow the flow of digesta through the tract. Bacterial cell walls are made primarily of peptidoglycan, which is hydrolyzed by the enzyme lysozyme. In an uneconomical match, the enzymatic and absorptive capacities would be in great excess relative to the typical load (i.e., the flow rate of primary nutrient) and/or retention time would be routinely in great excess in relation to reaction rates. The diffusive component of intestinal glucose absorption is mediated by the glucose-induced recruitment of GLUT2 to the brush-border membrane. Learning Objectives. Diet drives convergence in gut microbiome functions across mammalian phylogeny and within humans. Absorption refers to the transfer of compounds from the gut lumen across the gut wall to the body tissues, including the lymph or blood of vertebrates and hemolymph of arthropods. Adaptive variation in digestive enzyme activity with diet composition is crucial to the lifestyle of many animals. Reynolds DA, Rajendran VM, Binder HJ. Effects of diet quality on phenotypic flexibility of organ size and digestive function in Mongolian gerbils (. : Pyralidae). The https:// ensures that you are connecting to the A curious feature of the colonic rabbit lysozyme is that its pH optimum is very different from that of other lysozymes expressed in rabbits. Among animals that consume refractory food types there are multiple strategies. [SGLT1 expression has not been found to be influenced by cdx in mammals (405)]. Cai KH, Bennick A. Grajal A, Strahl SD, Parra R, Dominguez MG, Neher A. Foregut fermentation in the hoatzin, a neotropical leaf-eating bird. Marshall SDG, Gatehouse LN, Becher SA, Christeller JT, Gatehouse HS, Hurst MRH, Boucias DG, Jackson TA. It is a brush border enzyme that hydrolyzes monophosphate esters, but its physiological role in digestion has not been well understood. Niemann-Pick C1 Like 1 protein is critical for intestinal cholesterol absorption. Many insects, mammals, and birds respond by increasing secretion of proteolytic enzymes and, in the vertebrates, by increasing the size of the pancreas, which synthesizes many of the enzymes, often with the net effect of restoring digestive efficiency and growth rate. Learn. Several reviews are available regarding their interactions with SMs (299, 331, 412). Weiss SL, Lee EA, Diamond J. Voght SP, Fluegel ML, Andrews LA, Pallanck LJ. Meissner B, Boll M, Daniel H, Baumeister R. Deletion of the intestinal peptide transporter affects insulin and TOR signaling in Caenorhabditis elegans. Secretion of colonic isozyme of lisozyme in association with cecotrophy in rabbits. Evidence from digestive enzyme activities, gastrointestinal fermentation, and luminal nutrient concentrations. ), Polyphenolics (anthraquinone, proanthocyanidins and other tannins. Chen H, Pan YX, Wong EA, Webb KE. In contrast, absorption of 3-Omethyl-d-glucose did not differ significantly between the taxa. This means that the pig uterus has two large horns in addition to the body. (2)]. 11), and it used to be assumederroneouslythat cholesterol is taken up exclusively by simple diffusion. Knott KK, Barboza PS, Bowyer RT. Bergerson O, Wool D. The process of adaptation of flour beetles to new environments. The efflux of unhydrolyzed peptides across the basolateral membrane is mediated by peptide transporters that have not been identified at molecular level. Canavoso LE, Jouni ZE, Karnas KJ, Pennington JE, Wells MA. Onal U, Langdon C, Celik I. Ontogeny of the digestive tract of larval percula clownfish, Amphiprion percula (Lacepede 1802): A histological perspective. Many species respond to higher food intake by flexibly increasing digestive compartment size. In an another phylogenetically informed analysis, German et al. Karasov and colleagues measured total absorption (mediated and passive) of D-glucose or 3OMD-glucose and passive absorption of L-glucose in intact animals by a standard pharmacokinetic methodology, for example, references (78, 244, 278, 280). Fowler HG, Forti LC, Brandao CRF, Delabie JHC, Vasconcelos HL. In contrast, the anthraquinone, emodin, which tends to speed digesta through the gut of humans (137), appears to have the opposite effect on the frugivorous bird the Yellow-vented bulblul, and increases the birds apparent digestive efficiency on emodin-containing fruit (440). Chemicals from many of the major groupings of SMs (e.g., alkaloids, phenolics, and terpenoids) inhibit animals intrinsic mechanisms of breakdown of carbohydrates, fats, and proteins (Table 4). Tadmor-Melamed H, Markman S, Arieli A, Distl M, Wink M, Izhaki I. Many of these patterns are apparent in at least a dozen other species of mammals that have been studied, although in species such as carnivorous marine mammals and ruminants sucrase activity remains low (246), and in ruminants dramatic changes occur in GI tract structure postnatally [i.e., development of multichambered foregut (257, 258)] coordinated with changes in gene expression (97). Shifts during development in feeding versus nonfeeding or in dietary habits occur in diverse invertebrates, including lobsters (235) and insects (301), and digestive enzyme levels may change in correlation with changes in the major dietary substrates. Mazumdar-Leighton S, Broadway RM. Nutrition and ontogenetic development of the intestine. Most mammals and birds have a single gene copy that codes for lysozyme. Intestinal disaccharidases of young turkeys: Temporal development and influence of diet composition. Rumen-like methanogens identified from the crop of the folivorous South American bird, the hoatzin (Opisthocomus hoazin). Contributions of microbes in vertebrate gastrointestinal tract to production and conservation of nutrients. Some notable examples include evaluation of the glandular digestion path in lamellibranch bivalves that involves both intracellular digestion and extracellular digestion in the gut lumen (360), or compartmentalization imparted by the peritrophic envelope and enzyme recycling thought to occur in insects (34). Buddington RK, Chen JW, Diamond JM. Secondary metabolite emodin increases food assimilation efficiency of Yellow-vented Bulbuls (. The pig stomach is two to three times larger and the cardiac mucosa occupies a greater portion of the stomach compared to the human stomach. Some are very specific, for example, NAT6 and NAT8 in the distal midgut of mosquito Anopheles gambiae transport just aromatic amino acids (318, 319). Altmann SW, Davis HR, Jr, Zhu LJ, Yao X, Hoos LM, Tetzloff G, Iyer SP, Maguire M, Golovko A, Zeng M, Wang L, Murgolo N, Graziano MP. Sell JL, Koldovsky O, Reid BL. Lowry JB, McSweeney CS, Palmer B. Allometry of paracellular absorption in birds. Murray HM, Gallant JW, Johnson SC, Douglas SE. In some animals, the gut microbiota contributes directly to nutrition by the fermentative degradation of plant cell-wall polysaccharides. Kimmich GA, Randles J. Phloretin-like action of bioflavonoids on sugar accumulation capability of isolated intestinal cells. This issue has been explored particularly in relation to variation in the capacity of animal species with different diets to modulate their transporter activity. Skopec and Karasov (408) predicted that phloridzin would inhibit glucose absorption at the whole animal level when administered at ecological concentrations (they used 10 mmol/L), and that the effects would be more pronounced in nonflying mammals that rely on mediated pathway(s) for glucose absorption than birds that rely more on a nonmediated, paracellular pathway. Ruminants, colobine monkeys, and hoatzins have evolved independently a lysozyme that functions as a digestive enzyme [reviewed in reference (248)]. 5A) and maltase activity (Fig. Spiller HA, Winter MI, Weber JA, Krenzelok EP, Anderson II, Ryan MI. In mammals, a steep diffusion gradient across the apical membrane is generated by acyl-CoA:cholesterol acyltransferase (ACAT2)-mediated esterification of cholesterol in the enterocyte (Fig. The mechanism of chylomicron assembly is reviewed by reference (227). It can transport thousands of di- and tripeptides with low affinity and high capacity, but neither free amino acids nor tetrapeptides (106). The production of intrinsic cellulases by arthropods (insects), crustaceans (crayfish), and nematodes has been firmly established (463), but this capability is apparently absent from all vertebrates. German DP, Horn MH. The Digestive System in Mammal: Food, Form and Function. Common cutworms (Spodoptera litura; Lepidoptera), a highly polyphagous pest of subtropical and tropical crops, can be used to illustrate a pattern that is probably common (488). the contents by NLM or the National Institutes of Health. Caviedes-Vidal E, McWhorter TJ, Lavin SR, Chediack JG, Tracy CR, Karasov WH. Developmental changes in morphometry of the small intestine and jejunal sucrase activity during the first nine weeks of postnatal growth in pigs. Fish amylases and glucose transporters appear to be molecularly closely related to those in mammals and to have comparable characteristics (165, 269). In neonate rats, luminal or dietary carbohydrate does not induce glucose transport (which is contrary to the situation in adult rats), whereas in lambs dietary glucose is required for induction of glucose transport activity. Natural toxins are ubiquitous in foods and may influence key features such as digesta transit, enzymatic breakdown, microbial fermentation, and absorption. When rats reingest feces (coprophagy, or cecotrophy in rabbits), they digest and absorb labeled amino acid from those microbial proteins (Fig. It has been argued that pregastric fermentation chambers may have evolved in relation to functions other than cellulose degradation, for example to facilitate microbial detoxification of allelochemicals in ingested plant foods, and only subsequently became important in digestion of plant material (233). Based on arguments of the economy of nature (above), a number of patterns are predicted for animals adapted to particular diet features. Feldman DH, Harvey WR, Stevens BR. Warnecke F, Luginbuhl P, Ivanova N, Ghassemian M, Richardson TH, Stege JT, Cayouette M, McHardy AC, Djordjevic G, Aboushadi N, Sorek R, Tringe SG, Podar M, Martin HG, Kunin V, Dalevi D, Madejska J, Kirton E, Platt D, Szeto E, Salamov A, Barry K, Mikhailova N, Kyrpides NC, Matson EG, Ottesen EA, Zhang X, Hernandez M, Murillo C, Acosta LG, Rigoutsos I, Tamayo G, Green BD, Chang C, Rubin EM, Mathur EJ, Robertson DE, Hugenholtz P, Leadbetter JR. Metagenomic and functional analysis of hindgut microbiota of a wood-feeding higher termite. In another example, when larvae of bean weavils (Zabrotes subfasciatus) were fed seeds of Phaseolus vulgaris they secreted inducible isoforms of alpha-amylases that were insensitive to the alpha-amylase inhibitor that is found in the plant, whereas their constitutively produced alpha-amylase was inhibited by SMs in the plant [reference (29); see also references (29, 403)]. Dietary protein and energy as determinants of food quality: Trophic strategies compared. Brzek P, Lessner KM, Caviedes-Vidal E, Karasov WH. Hindgut fermenting animals may also digest bacteria when they reingest their feces (coprophagy/cecotrophy). Binding of phlorizin to the C-terminal loop 13 of the Na+/glucose cotransporter does not depend on the 560608 disulfide bond. Sklan D, Noy Y. Functional development and intestinal absorption in the young poult. L-glucose absorption in house sparrows (Passer domesticus) is nonmediated. In: Chivers DJ, Langer P, editors. Fermentation and gstrointestinal microorganisms in fishes. 17 C), suggesting that it may have initiated transcription of SI (405). 9) (206). Honma K, Mochizuki K, Goda T. Carbohydrate/fat ratio in the diet alters histone acetylation on the sucrase-isomaltase gene and its expression in mouse small intestine. Fully reversible phenotypic plasticity of digestive physiology in young house sparrows: Lack of long-term effect of diet composition. Chediack JG, Caviedes-Vidal E, Karasov WH. government site. A model of digestion modulation in grasshoppers. Bifano TD, Alegria TG, Terra WR. The enzymes important for digestion can be clarified based on cDNA sequence (e.g., particular catalytic motifs), tissue localization (by fluorescent in situ hybridization of mRNA and immunohistochemistry of protein), and developmental and induced expression (e.g., during feeding vs. nonfeeding stages) (487, 488). The taxon richness of the gut microbiota, usually identified by 16S rRNA gene sequencing, is typically an order of magnitude greater in vertebrates than invertebrates, and the interspecific variation in microbial composition is strongly influenced by diet. In this section, two aspects of nutrient absorption are addressed: the modes of transport of the major classes of organic solutes and variation in nutrient absorption among animal taxa, in relation to nutritional habits and phylogeny and its mechanistic basis. Kohl KD, Brzek P, Caviedes-Vidal E, Karasov WH. Purification and partial characterization of a midgut membrane-bound alpha-glucosidase from. Digesta passage, digestibility and behavior in captive gorillas under two dietary regimens. (366) showed that lipopolysaccharide (LPS), a major component of the bacterial outer membrane, acts as a substrate at physiologically relevant pH. The assimilation of bacterial protein by herbivorous birds is perplexing because birds do not seem to have spatial separation of culturing and digestion of microbes. Gilbert ER, Williams PM, Ray WK, Li HF, Emmerson DA, Wong EA, Webb KE. For example, in response to high dietary supply of sugars, the expression of genes encoding the transporters SGLT1 (for glucose) and GLUT5 (for fructose) is increased. These enzymes are active against the sulfated polysaccharides in Porphyra seaweeds that form a regular part of the typical Japanese, but not North American, diet. Verri T, Kottra G, Romano A, Tiso N, Peric M, Maffia M, Boll M, Argenton F, Daniel H, Storelli C. Molecular and functional characterisation of the zebrafish (Danio rerio) PEPT1-type peptide transporter. Santo Domingo JW, Kaufman MG, Klug MJ, Holben WE, Harris D, Tiedge JM. (A) Functional groups of bacteria (SRBs, sulfate-reducing bacteria). It is not known whether such genetic or phenotypic adaptive response to dietary glycosides occurs in a vertebrate species. Although the contribution of the various tight junction proteins to the restriction of movement between epithelial cells is not fully understood, there is growing evidence that: (i) the claudins (a family of membrane proteins spanning the tight junction) play a crucial role in the pore pathway, with individual family members forming cation- or anion-selective pores; (ii) two further tight junction proteins, occludin and zona occludens-1, are important in the leak pathway; and (iii) various intracellular and extracellular signals mediate cross-talk between the two pathways, resulting in dynamic regulation of flux of different classes of compounds by the paracellular route. Development of intestinal transport function in mammals. Baker JE, Lum PTM, Halliday WR. Fructose is transported principally via the facilitative transporter GLUT5 (126). Their functions include communication, attraction, or in defense against herbivores, predators, pathogens, and competitors (202). (B) Major bacterial taxa responsible for degradation of starch and fructan-carbohydrates. Examples of organ systems include the cardiovascular, respiratory, and digestive . Multiple factors beyond the biochemical capabilities of the microbiota determine the nutritional significance of microbial fermentation for an animal. Froystad MK, Lilleeng E, Sundby A, Krogdahl A. Cloning and characterization of alpha-amylase from Atlantic salmon (, Fujita A, Shimizu I, Abe T. Distribution of lysozyme and protease, and amino acid concentration in the guts of a wood-feeding termite. Although within any single food category, there can be tremendous variation, some generalities emerge. This mode of regulation both maximizes the digestibility of substrates and minimizes the cost of synthesizing excess enzyme when the substrate is at low levels. Penry DL. Zhan QL, Zheng SC, Feng QL, Liu L. A midgut-specific chymotrypsin cDNA (SLCTP1) from, Zhang C, Zhou DH, Zheng SC, Liu L, Tao S, Yang L, Hu SN, Feng QL. Jakobsson HE, Jernberg C, Andersson AF, Sjolund-Karlsson M, Jansson JK, Engstrand L. Short-term antibiotic treatment has differing long-term impacts on the human throat and gut microbiome. Hepatocyte nuclear factor-1 alpha, GATA-4, and caudal related homeodomain protein Cdx2 interact functionally to modulate intestinal gene transcription. The liver, pancreas, and gallbladder are the solid organs of the digestive system. For example, metagenomic analyses have identified more than 700 candidate glucohydrolase genes of bacterial origin in the hindgut paunch of Nasutitermes termites, most of which have predicted capacity to degrade cellulose and xylans (462), and a remarkable 27,755 putative carbohydrate-active genes have been detected in the metagenome of the cow rumen contents, most of which are bacterial in origin, have less than 75% sequence identity with previously described genes, and many of which are likely active against cellulose (210).

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