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Intestinal Calcium Absorption

Kannikar Wongdee, Krittikan Chanpaisaeng, Jarinthorn Teerapornpuntakit, Narattaphol Charoenphandhu

10.1002/cphy.c200014

Source: Volume 11, Issue 3, July 2021

Published online: May 2021

Full Article on Wiley Online Library



Abstract

In this article, we focus on mammalian calcium absorption across the intestinal epithelium in normal physiology. Intestinal calcium transport is essential for supplying calcium for metabolism and bone mineralization. Dietary calcium is transported across the mucosal epithelia via saturable transcellular and nonsaturable paracellular pathways, both of which are under the regulation of 1,25‐dihydroxyvitamin D3 and several other endocrine and paracrine factors, such as parathyroid hormone, prolactin, 17β‐estradiol, calcitonin, and fibroblast growth factor‐23. Calcium absorption occurs in several segments of the small and large intestine with varying rates and capacities. Segmental heterogeneity also includes differential expression of calcium transporters/carriers (e.g., transient receptor potential cation channel and calbindin‐D9k) and the presence of favorable factors (e.g., pH, luminal contents, and gut motility). Other proteins and transporters (e.g., plasma membrane vitamin D receptor and voltage‐dependent calcium channels), as well as vesicular calcium transport that probably contributes to intestinal calcium absorption, are also discussed. © 2021 American Physiological Society. Compr Physiol 11:2047‐2073, 2021.

Figure 1. Figure 1. Segmental heterogeneity of calcium absorption. Key features including magnitude of net absorptive calcium flux rate (Jnet = Jmucosa‐to‐serosa − Jserosa‐to‐mucosa) and percentage of total absorbed calcium under normal (calcium‐ and vitamin D‐replete) conditions are presented for each intestinal segment. The number of “+” signs represents the amount of net calcium flux compared to cecum which has the highest net flux marked with “++++”.
Figure 2. Figure 2. Intestinal segment‐specific sojourn time, calcium solubility, and expression profile of genes/proteins related to calcium transport. The number of “+” signifies the relative abundance of a feature for a given intestinal segment. A “–” indicates that an mRNA expression is absent in a segment. The sojourn time in the table represents time of chyme. The lower lines of each segment (red text) represent distribution of mRNA expression of genes related to calcium transport in female rats refer to our previous study 219. N/A, not available; S, sexual disparity in the expression profile; Cldn, claudin; PMCA1b, plasma membrane Ca2+ ATPase‐1b; TRPV6, transient receptor potential cation channel, subfamily V, member 6; VDR, vitamin D receptor.
Figure 3. Figure 3. Diagrams of intestinal villi. (A) Longitudinal section of villous showing sites of net calcium absorption at villous (a) and net calcium secretion at crypts of intestine (b). The heterogeneity of calbindin‐D9k (S100 calcium‐binding protein G, S100G) expression and the primary direction of calcium flux are presented along the crypt‐villous axis. (B) Transverse section of villous tip featuring transcellular (c) and paracellular calcium absorption (d), secretion (e), and the existence of unstirred water layer and acid microclimate (f).
Figure 4. Figure 4. Mechanism of transcellular calcium transport across the enterocyte. Calcium is transported across apical membrane through transient receptor potential cation channel, subfamily V, member 6 (TRPV6), and, to a lesser extent, TRPV5, facilitated diffusion through the cytosol, and active basolateral extrusion by the plasma membrane Ca2+ ATPase‐1b (PMCA1b) and other transporters [e.g., Na+/Ca2+ exchanger 1 (NCX1) and K+‐dependent Na+/Ca2+ exchanger (NCKX)]. Some inhibitors of each calcium transporter are also indicated. Calcium‐binding proteins including calbindin‐D9k (CaBP‐9k) and calmodulin (CAM) may play a role in a buffering system to prevent intracellular calcium overload as well as translocate calcium ions from the mucosa to the serosa. The stoichiometry for NCX1 is 4 Na+ to 1‐2 Ca2+ moved per transport cycle; however, an average stoichiometry is 3 Na+:1 Ca2+. The stoichiometry of NKA is 3 Na+:2 K+. TRPV6, NKA, and NCX1 activities are inhibited by ruthenium red, ouabain, and KB‐R7943, respectively. PMCA1b activity is inhibited by trifluoperazine and vanadate.
Figure 5. Figure 5. Effect of apical membrane potential changes on transcellular intestinal calcium absorption. During glucose uptake via Na+‐dependent glucose transporter (SGLT)‐1, sodium entry induces slight depolarization of the apical membrane, thereby triggering Cav1.3 opening at slight depolarized potential of approximately −40 mV as compared with −47 mV resting potential of the apical membrane. Cav1.3 activity is inhibited by dihydropyridine. PMCA1b, plasma membrane Ca2+ ATPase‐1b; NCX1, Na+/Ca2+ exchanger 1; NKA, Na+/K+‐ATPase; calbindin‐D9k (S100 calcium‐binding protein G, S100G); CAM, calmodulin.
Figure 6. Figure 6. Model of vesicular and reticular transport of calcium. Diagram shows vesicular calcium transport and calcium tunneling through the meshwork of endoplasmic reticulum (ER). Calcium enters the cell through TRPV6 or other calcium channels. Intracellular calcium pool (Cai) in the apical compartment is moved into the vesicles or pumped into ER lumen by sarco‐endoplasmic reticulum Ca2+‐ATPase (SERCA). Calcium diffuses along the ER tunnel to the basolateral side and released into the cytoplasm by 1,4,5‐trisphosphate receptor (IP3R) and ryanodine receptor (RyR) and extrudes by the plasma membrane Ca2+ ATPase‐1b (PMCA1b). In cellular calcium depleted condition, a calcium sensor stromal interacting molecule 1 (STIM1) is translocated to the apical membrane to induce electrogenic calcium influx. The pathways with red texts (i.e., STIM1, SERCA, ER, IP3R, and RyR) are hypothetical from indirect evidence and need more investigations to confirm their physiological significance. TRPV6, transient receptor potential cation channel, subfamily V, member 6.
Figure 7. Figure 7. Paracellular intestinal calcium transport. Although calcium concentration gradient is a driving force for paracellular calcium transport, the tight junction (i.e., claudins, occludin, ZO‐1) acts as a barrier to restricts paracellular ion movement in a charge‐ and size‐selective manner. Calcium and small water‐soluble molecules move along the stream of water in apical to basolateral direction (solvent drag‐induced calcium absorption). Elevation of NKA activity increases the driving force for the solvent drag‐induced calcium transport. ZO‐1, zonula occludens‐1.
Figure 8. Figure 8. Systemic control of mammalian calcium homeostasis by three organs (i.e., bone, kidney, and intestine). The details are fully explained in the section titled “Systemic Controls Mucosal Calcium Absorption”. Cyp27B1, 25‐hydroxyvitamin D3 1α‐hydroxylase; PTH, parathyroid hormone.
Figure 9. Figure 9. Extrinsic (humoral and neural) and intrinsic (luminal) regulators of intestinal calcium absorption. The luminal and endocrine factors that act as stimulator and inhibitor on intestinal calcium absorption showing in the green and orange boxes, respectively. Although the presented luminal and endocrine factors are known to modulate calcium absorption, the involvement of neurocrine factors (marked as italic text), or factors released from neuroendocrine cells in the enteric nervous system is in the emerging area of interest and needs more investigations to confirm the physiological significance. Cldn, claudin; FGF‐23, fibroblast growth factor‐23; GH, growth hormone; IGF‐1, insulin‐like growth factor‐1; SCFAs, short‐chain fatty acids; TTX, tetrodotoxin; VIP, vasoactive intestinal peptide; NCX1, Na+/Ca2+ exchanger 1; NKA, Na+/K+‐ATPase; PMCA1b, plasma membrane Ca2+ ATPase‐1b; calbindin‐D9k (S100 calcium‐binding protein G, S100G); TRPV, transient receptor potential cation channel, subfamily V.
Figure 10. Figure 10. The emerging topic of intestinal calcium absorption. (A) A schematic diagram shows possible mechanism of fibroblast growth factor (FGF)‐23 and calcium‐sensing receptor (CaSR) on local negative feedback regulation of calcium absorption. (B) A schematic diagram showing a possible mechanism of how short‐chain fatty acids (SCFAs) enhances transcellular calcium transport in the intestine. Hypothetical pathways or pathways with indirect evidence are presented as dashed lines. NCX1, Na+/Ca2+ exchanger 1; NKA, Na+/K+‐ATPase; PMCA1b, plasma membrane Ca2+ ATPase‐1b; calbindin‐D9k (S100 calcium‐binding protein G, S100G); TRPV, transient receptor potential cation channel, subfamily V.


Figure 1. Segmental heterogeneity of calcium absorption. Key features including magnitude of net absorptive calcium flux rate (Jnet = Jmucosa‐to‐serosa − Jserosa‐to‐mucosa) and percentage of total absorbed calcium under normal (calcium‐ and vitamin D‐replete) conditions are presented for each intestinal segment. The number of “+” signs represents the amount of net calcium flux compared to cecum which has the highest net flux marked with “++++”.


Figure 2. Intestinal segment‐specific sojourn time, calcium solubility, and expression profile of genes/proteins related to calcium transport. The number of “+” signifies the relative abundance of a feature for a given intestinal segment. A “–” indicates that an mRNA expression is absent in a segment. The sojourn time in the table represents time of chyme. The lower lines of each segment (red text) represent distribution of mRNA expression of genes related to calcium transport in female rats refer to our previous study 219. N/A, not available; S, sexual disparity in the expression profile; Cldn, claudin; PMCA1b, plasma membrane Ca2+ ATPase‐1b; TRPV6, transient receptor potential cation channel, subfamily V, member 6; VDR, vitamin D receptor.


Figure 3. Diagrams of intestinal villi. (A) Longitudinal section of villous showing sites of net calcium absorption at villous (a) and net calcium secretion at crypts of intestine (b). The heterogeneity of calbindin‐D9k (S100 calcium‐binding protein G, S100G) expression and the primary direction of calcium flux are presented along the crypt‐villous axis. (B) Transverse section of villous tip featuring transcellular (c) and paracellular calcium absorption (d), secretion (e), and the existence of unstirred water layer and acid microclimate (f).


Figure 4. Mechanism of transcellular calcium transport across the enterocyte. Calcium is transported across apical membrane through transient receptor potential cation channel, subfamily V, member 6 (TRPV6), and, to a lesser extent, TRPV5, facilitated diffusion through the cytosol, and active basolateral extrusion by the plasma membrane Ca2+ ATPase‐1b (PMCA1b) and other transporters [e.g., Na+/Ca2+ exchanger 1 (NCX1) and K+‐dependent Na+/Ca2+ exchanger (NCKX)]. Some inhibitors of each calcium transporter are also indicated. Calcium‐binding proteins including calbindin‐D9k (CaBP‐9k) and calmodulin (CAM) may play a role in a buffering system to prevent intracellular calcium overload as well as translocate calcium ions from the mucosa to the serosa. The stoichiometry for NCX1 is 4 Na+ to 1‐2 Ca2+ moved per transport cycle; however, an average stoichiometry is 3 Na+:1 Ca2+. The stoichiometry of NKA is 3 Na+:2 K+. TRPV6, NKA, and NCX1 activities are inhibited by ruthenium red, ouabain, and KB‐R7943, respectively. PMCA1b activity is inhibited by trifluoperazine and vanadate.


Figure 5. Effect of apical membrane potential changes on transcellular intestinal calcium absorption. During glucose uptake via Na+‐dependent glucose transporter (SGLT)‐1, sodium entry induces slight depolarization of the apical membrane, thereby triggering Cav1.3 opening at slight depolarized potential of approximately −40 mV as compared with −47 mV resting potential of the apical membrane. Cav1.3 activity is inhibited by dihydropyridine. PMCA1b, plasma membrane Ca2+ ATPase‐1b; NCX1, Na+/Ca2+ exchanger 1; NKA, Na+/K+‐ATPase; calbindin‐D9k (S100 calcium‐binding protein G, S100G); CAM, calmodulin.


Figure 6. Model of vesicular and reticular transport of calcium. Diagram shows vesicular calcium transport and calcium tunneling through the meshwork of endoplasmic reticulum (ER). Calcium enters the cell through TRPV6 or other calcium channels. Intracellular calcium pool (Cai) in the apical compartment is moved into the vesicles or pumped into ER lumen by sarco‐endoplasmic reticulum Ca2+‐ATPase (SERCA). Calcium diffuses along the ER tunnel to the basolateral side and released into the cytoplasm by 1,4,5‐trisphosphate receptor (IP3R) and ryanodine receptor (RyR) and extrudes by the plasma membrane Ca2+ ATPase‐1b (PMCA1b). In cellular calcium depleted condition, a calcium sensor stromal interacting molecule 1 (STIM1) is translocated to the apical membrane to induce electrogenic calcium influx. The pathways with red texts (i.e., STIM1, SERCA, ER, IP3R, and RyR) are hypothetical from indirect evidence and need more investigations to confirm their physiological significance. TRPV6, transient receptor potential cation channel, subfamily V, member 6.


Figure 7. Paracellular intestinal calcium transport. Although calcium concentration gradient is a driving force for paracellular calcium transport, the tight junction (i.e., claudins, occludin, ZO‐1) acts as a barrier to restricts paracellular ion movement in a charge‐ and size‐selective manner. Calcium and small water‐soluble molecules move along the stream of water in apical to basolateral direction (solvent drag‐induced calcium absorption). Elevation of NKA activity increases the driving force for the solvent drag‐induced calcium transport. ZO‐1, zonula occludens‐1.


Figure 8. Systemic control of mammalian calcium homeostasis by three organs (i.e., bone, kidney, and intestine). The details are fully explained in the section titled “Systemic Controls Mucosal Calcium Absorption”. Cyp27B1, 25‐hydroxyvitamin D3 1α‐hydroxylase; PTH, parathyroid hormone.


Figure 9. Extrinsic (humoral and neural) and intrinsic (luminal) regulators of intestinal calcium absorption. The luminal and endocrine factors that act as stimulator and inhibitor on intestinal calcium absorption showing in the green and orange boxes, respectively. Although the presented luminal and endocrine factors are known to modulate calcium absorption, the involvement of neurocrine factors (marked as italic text), or factors released from neuroendocrine cells in the enteric nervous system is in the emerging area of interest and needs more investigations to confirm the physiological significance. Cldn, claudin; FGF‐23, fibroblast growth factor‐23; GH, growth hormone; IGF‐1, insulin‐like growth factor‐1; SCFAs, short‐chain fatty acids; TTX, tetrodotoxin; VIP, vasoactive intestinal peptide; NCX1, Na+/Ca2+ exchanger 1; NKA, Na+/K+‐ATPase; PMCA1b, plasma membrane Ca2+ ATPase‐1b; calbindin‐D9k (S100 calcium‐binding protein G, S100G); TRPV, transient receptor potential cation channel, subfamily V.


Figure 10. The emerging topic of intestinal calcium absorption. (A) A schematic diagram shows possible mechanism of fibroblast growth factor (FGF)‐23 and calcium‐sensing receptor (CaSR) on local negative feedback regulation of calcium absorption. (B) A schematic diagram showing a possible mechanism of how short‐chain fatty acids (SCFAs) enhances transcellular calcium transport in the intestine. Hypothetical pathways or pathways with indirect evidence are presented as dashed lines. NCX1, Na+/Ca2+ exchanger 1; NKA, Na+/K+‐ATPase; PMCA1b, plasma membrane Ca2+ ATPase‐1b; calbindin‐D9k (S100 calcium‐binding protein G, S100G); TRPV, transient receptor potential cation channel, subfamily V.
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Article Title: Intestinal Calcium Absorption
 

How to Cite

Kannikar Wongdee, Krittikan Chanpaisaeng, Jarinthorn Teerapornpuntakit, Narattaphol Charoenphandhu. Intestinal Calcium Absorption. Compr Physiol 2021, 11: 2047-2073. doi: 10.1002/cphy.c200014