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Iron in Infection and Immunity
Open accessAbstract
Iron is an essential nutrient for both humans and pathogenic microbes. Because of its ability to exist in one of two oxidation states, iron is an ideal redox catalyst for diverse cellular processes including respiration and DNA replication. However, the redox potential of iron also contributes to its toxicity; thus, iron concentration and distribution must be carefully controlled. Given the absolute requirement for iron by virtually all human pathogens, an important facet of the innate immune system is to limit iron availability to invading microbes in a process termed nutritional immunity. Successful human pathogens must therefore possess mechanisms to circumvent nutritional immunity in order to cause disease. In this review, we discuss regulation of iron metabolism in the setting of infection and delineate strategies used by human pathogens to overcome iron-withholding defenses.
Overview
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Written from the paper’s full text.
TL;DR
This review argues that because nearly all human pathogens require iron, withholding iron from microbes (nutritional immunity) is a central innate immune strategy, and successful pathogens must evolve countermeasures to cause disease. It surveys host iron homeostasis, microbial acquisition and detoxification strategies across bacteria, fungi, and parasites, and therapeutic approaches that target pathogen iron metabolism.
Key findings
- Human iron metabolism is highly conserving: adults hold roughly 4–5 g of total body iron while losing only about 0.5–1 mg daily, and healthy transferrin is typically under 50% iron-saturated, leaving little free extracellular iron for microbes.
- Infection triggers hypoferremia, orchestrated largely by hepcidin (which drives ferroportin internalization and degradation) plus hepcidin-independent cytokine effects; local sequestration adds lactoferrin, Nramp1, and siderocalin/lipocalin-2.
- Pathogens use five main acquisition routes — manganese substitution (e.g., *Borrelia burgdorferi*), siderophores (binding constants exceeding 10^50 versus transferrin's ~10^36), heme uptake systems, transferrin/lactoferrin receptors, and ferric/ferrous transporters — and some produce "stealth siderophores" that evade siderocalin.
- Iron overload states such as hereditary hemochromatosis and thalassemia raise infection susceptibility (e.g., *Yersinia* spp., *Listeria monocytogenes*, *Vibrio vulnificus*), though hemochromatosis monocytes are reportedly less permissive to *M. tuberculosis*.
- Therapeutic avenues include siderophore-biosynthesis inhibitors (e.g., p-aminosalicylic acid), sideromycin conjugates (albomycin, MC-1), and vaccines against iron-acquisition proteins (IsdA/IsdB, uropathogenic *E. coli* iron receptors, hookworm Na-APR-1).
Methods
Narrative review; no primary data or systematic search strategy is described. Evidence is synthesized from cited primary literature spanning human iron physiology, animal and in vitro infection models, and clinical observations (e.g., the 1940s canine *S. aureus* hypoferremia experiments, mouse models, and human vaccine trial outcomes). Sections are organized by host homeostasis, bacterial (extracellular and intracellular), fungal, and parasitic iron handling, then therapeutics.
Limitations
The paper states several open questions: which redundant microbial iron-acquisition components matter most for pathogenesis, which are best suited to vaccine construction, whether the hepcidin axis can be therapeutically manipulated for tolerance, and whether mammalian siderophores function in nutritional immunity. It notes that the terminated *S. aureus* IsdB-containing human vaccine trial shows animal protection may not predict human efficacy. Evident from the text: as a review, it offers no new experimental data, and several claims rest on single or dated sources (e.g., textbook citations for epidemiologic figures such as hookworm and malaria burden).
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In the paper
“Acquisition of heme by the Isd system is critical for S. aureus pathogenesis (Cassat and Skaar, 2012).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Cassat and Skaar, 2012 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In addition to mitigating toxicity associated with hypo- or hyperferremia, regulation of iron distribution serves as an innate immune mechanism against invading pathogens.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Patients with iron overload from hemochromatosis and thalassemia are more susceptible to infection with Yersinia spp., Listeria monocytogenes, and Vibrio vulnificus, among others (Long et al., 2003).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ic. Disorders of iron homeostasis that enhance susceptibility to infectious diseases
What the sources say
- Not checked yetcites Long et al., 2003
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In the paper
“Nramp1 (natural resistance associated macrophage protein-1) is a divalent metal ion transporter that has been proposed to reduce iron content in early endosomes, thereby limiting availability to intracellular pathogens in this compartment (Jabado et al., 2000; Vidal et al., 1995).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
Strength:hedged· not yet calibratedCausal:conditional· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesThe paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“However, hemolytic pathogens must subsequently compete with haptoglobin and hemopexin, host glycoproteins that scavenge liberated hemoglobin and heme, respectively (Figure 1D).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Inside the macrophage, Mtb disrupts phagolysosomal maturation, an iron-dependent process critical for pathogen survival (Olakanmi et al., 2000).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIb. Iron acquisition by intracellular bacterial pathogens
What the sources say
- Not checked yetcites Olakanmi et al., 2000 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIn the paper
“Siderophores can therefore effectively outcompete host transferrin, which binds iron with an association constant of approximately 1036.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Bullen JJ, 1999
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIn the paper
“Chelation therapy with desferrioxamine may therefore increase the risk of infection in patients with iron overload (Kim et al., 2007).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Kim et al., 2007 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The importance of strict regulation of iron metabolism is underscored by the pathologic consequences of iron overload syndromes. Individuals suffering from iron overload not only experience cellular damage from adverse redox chemistry of free iron, but also are at enhanced risk of infection.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ic. Disorders of iron homeostasis that enhance susceptibility to infectious diseases
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesA conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Hepcidin is a major orchestrator of the hypoferremic response to infection.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passes
- It may state two separate claims as one. 39
- Probably not a substantial claim. 7
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Because of its ability to exist in one of two oxidation states, iron is an ideal redox catalyst for diverse cellular processes including respiration and DNA replication.”
- Also stated in: § Ia. Human iron homeostasis · plain
“The biologic utility of iron resides in its ability to cycle between two oxidation states: ferrous (Fe2+) or ferric (Fe3+). Iron can thus serve as a redox catalyst, accepting or donating electrons.”
§ Iron in Infection and Immunity > Abstract
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
- Also stated in: § Ia. Human iron homeostasis · plain
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Iron is an essential nutrient for both humans and pathogenic microbes.”
- Also stated in: § Ia. Human iron homeostasis · plain
“Iron is an essential nutrient for humans, with critical functions in many cellular processes.”
§ Iron in Infection and Immunity > Abstract
Load-bearingStrength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
- Also stated in: § Ia. Human iron homeostasis · plain
In the paper
“For example, V. vulnificus, S. aureus, and Y. enterocolitica, among others, can utilize desferrioxamine B as an iron source.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Kim et al., 2007 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Neisseria meningitidis expresses both transferrin and lactoferrin uptake systems. Consistent with its role as an obligate human pathogen, N. meningitidis preferentially utilizes human transferrin (Schryvers and Morris, 1988).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Schryvers and Morris, 1988 source
Strength:plain· not yet calibratedCausal:correlational· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Moreover, unlike transferrin, lactoferrin maintains iron-binding capacity at low pH and therefore may be a more effective scavenger in acidotic infectious foci (Baker and Baker, 2012).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
- Not checked yetcites Baker and Baker, 2012 source
Strength:hedged· not yet calibratedCausal:conditional· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Additionally, the specific (“secondary”) granules of neutrophils contain lactoferrin, which is released at infectious sites in response to cytokines (Masson et al., 1969).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
- Not checked yetcites Masson et al., 1969 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“In healthy individuals, nearly all iron released into plasma is bound to transferrin, limiting iron-catalyzed free radical production and facilitating transport to target cells.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ia. Human iron homeostasis
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“In fact, hepcidin was initially characterized as an antimicrobial peptide in human urine and blood ultrafiltrate (Krause et al., 2000; Park et al., 2001).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“First, the majority of iron in humans is sequestered intracellularly, complexed within hemoglobin inside erythrocytes.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Increases in total body iron stores trigger the production of hepcidin, which subsequently induces the internalization and degradation of ferroportin (Nemeth et al., 2004).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ia. Human iron homeostasis
What the sources say
- Not checked yetcites Nemeth et al., 2004 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“As most iron in humans is contained within hemoglobin, many pathogens have developed mechanisms to liberate hemoglobin from erythrocytes and utilize heme as a nutrient.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Hepcidin release from the liver is stimulated by pro-inflammatory cytokines, TLR activation, and induction of the endoplasmic reticulum unfolded protein response (Figure 2A) (Drakesmith and Prentice, 2012).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
- Not checked yetcites Drakesmith and Prentice, 2012 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Hepcidin, a peptide hormone produced in the liver, post-translationally regulates ferroportin and thus controls entry of iron into the plasma after enterocyte absorption.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ia. Human iron homeostasis
What the sources say
- Not checked yetcites Nemeth et al., 2004 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Produced by a wide range of Gram-positive and Gram-negative human pathogens, siderophores are small ferric iron chelators capable of binding iron with association constants that can exceed 1050 (Bullen JJ, 1999).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Bullen JJ, 1999
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“P. aeruginosa expresses both a direct uptake system and hemophore-mediated system for heme acquisition.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“The hypoferremia of infection was documented in seminal studies by2 Cartwright et al. in the 1940s, who noted a precipitous drop in plasma iron levels upon intramuscular inoculation of canines with Staphylococcus aureus.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
- Not checked yetcites Cartwright et al., 1946 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The hypoferremic response to infection is also mediated by hepcidin-independent mechanisms. Cytokines such as interferon-gamma, tumor necrosis factor-alpha, IL-1, and IL-6 modulate iron metabolism to further strengthen iron-withholding defenses (Figure 2B – C) (Nairz et al., 2010; Weiss, 2005).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“They are synthesized under conditions of low iron availability and secreted into the extracellular environment, where they bind ferric iron and facilitate its use by microbial cells (Figure 3A–3B).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“One mechanism, used by Borrelia burgdorferi, is to utilize manganese instead of iron in metalloenzymes, thereby eliminating the need for iron acquisition systems (Posey and Gherardini, 2000).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
What the sources say
- Not checked yetcites Posey and Gherardini, 2000 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Extracellular iron is bound with high affinity by transferrin, which in healthy individuals is typically less than 50% saturated with iron.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
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The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Lactoferrin is a host glycoprotein that, like transferrin, binds free iron with high affinity. Mucosal secretions contain high concentrations of lactoferrin, comprising a constitutive mechanism for iron limitation at mucosal surfaces.”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Siderocalin, also known as lipocalin-2 (Lcn2) or neutrophil gelatinase-associated lipocalin, is expressed during the acute phase response to infection and sequesters siderophores (Figure 2D).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Radiolabeling studies demonstrated that Mtb acquires iron either from transferrin as it cycles through the endocytic pathway, or from cytoplasmic iron stores (Olakanmi et al., 2002).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIb. Iron acquisition by intracellular bacterial pathogens
What the sources say
- Not checked yetcites Olakanmi et al., 2002 source
Strength:boosted· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Polymorphisms in Nramp1 have been found to correlate with enhanced human susceptibility to Mycobacterium tuberculosis (Mtb), suggesting that intraphagosomal iron limitation might be an important innate immune defense (van Crevel et al., 2009).”
§ Iron in Infection and Immunity > I. Mechanisms of iron homeostasis in human health and disease > Ib. Iron limitation as an innate immune defense
What the sources say
- Not checked yetcites van Crevel et al., 2009 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Similar to predominantly extracellular pathogens, pathogenic mycobacteria express siderophores known as mycobactins, the biosynthesis of which has been elegantly investigated using lipidomics profiling (Madigan et al., 2012). Disruption of mycobactin synthesis impairs growth within macrophages, suggesting that siderophore production is an important iron acquisition strategy for Mtb (De Voss et al.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIb. Iron acquisition by intracellular bacterial pathogens
What the sources say
- Not checked yetcites De Voss et al., 2000 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Siderocalin was initially characterized by its ability to bind the E. coli siderophore enterochelin, inhibiting its growth.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIa. Iron acquisition by predominantly extracellular bacterial pathogens
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Human hookworms infect greater than 700 million people worldwide and are a leading cause of iron deficiency anemia in the developing world (Long et al., 2003).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
- Not checked yetcites Long et al., 2003
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“After erythrocyte rupture, hemozoin is released into the circulation and ingested by phagocytes, where it has profound immunomodulatory effects (Hanscheid et al., 2007; Shio et al., 2010).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
Strength:boosted· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Human hookworms infect greater than 700 million people worldwide and are a leading cause of iron deficiency anemia in the developing world (Long et al., 2003).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
- Not checked yetcites Long et al., 2003
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Similar to human hookworms, P. falciparum also uses host hemoglobin as a nutrient source, and has evolved a distinct mechanism for protecting against iron toxicity.”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
- Rests on an assumption: Malaria is caused by one of five species of Plasmodium, with P. falciparum being the most virulent.
- Rests on an assumption: Hemoglobin is ingested by intra-erythrocytic Plasmodia, transported to a food vacuole, and degraded to nutrient amino acids by a proteolytic cascade.
In the paper
“Therefore, although hereditary hemochromatosis generally enhances susceptibility to infectious disease, certain subtypes may confer a selective advantage against intracellular pathogens.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIb. Iron acquisition by intracellular bacterial pathogens
What the sources say
- Not checked yetcites Olakanmi et al., 2002 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Lcn2 plays a critical role in control of parasitemia through suppression of reticulocytosis, enhancement of macrophage function, and modulation of adaptive immune responses (Zhao et al., 2012).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
- Not checked yetcites Zhao et al., 2012 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
- Rests on an assumption: Lcn2 is abundantly produced during malaria.
In the paper
“For example, disruption of Mtb ferritin biosynthesis limits resistance to oxidative stress, enhances susceptibility to antibiotics, and decreases bacterial survival in a chronic infection model (Pandey and Rodriguez, 2012).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIc. Iron detoxification strategies in bacterial pathogens
What the sources say
- Not checked yetcites Pandey and Rodriguez, 2012 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“HssRS senses heme exposure, increasing expression of hrtAB, which results in alleviation of heme toxicity.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIc. Iron detoxification strategies in bacterial pathogens
What the sources say
- Not checked yetcites Stauff et al., 2007 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Interestingly, patients with hereditary hemochromatosis have reduced intracellular iron levels, and thus monocytes derived from such individuals are less permissive to Mtb replication (Olakanmi et al., 2002).”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIb. Iron acquisition by intracellular bacterial pathogens
What the sources say
- Not checked yetcites Olakanmi et al., 2002 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Up to two-thirds of the world’s population are exposed to malaria annually, of which over 240 million will contract the disease and up to 2 million will die (Long et al., 2003).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
- Not checked yetcites Long et al., 2003
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Both antimicrobial compounds and vaccines targeting pathogen iron homeostasis are currently being developed.”
§ Iron in Infection and Immunity > V. Targeting microbial iron homeostasis for the treatment of infectious diseases
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
- Rests on an assumption: The vast majority of human pathogens require iron to sustain growth and colonize humans.
In the paper
“Additionally, ferroportin rapidly localizes to Mtb-infected phagolysosomes, resulting in iron efflux from this cellular compartment.”
§ Iron in Infection and Immunity > II. Iron acquisition by pathogenic microbes > IIb. Iron acquisition by intracellular bacterial pathogens
What the sources say
- Not checked yetcites Van Zandt et al., 2008 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Up to two-thirds of the world’s population are exposed to malaria annually, of which over 240 million will contract the disease and up to 2 million will die (Long et al., 2003).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
- Not checked yetcites Long et al., 2003
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Pathogenic fungi, like bacterial pathogens, must obtain iron from host tissues and regulate iron homeostasis to avoid toxicity.”
§ Iron in Infection and Immunity > III. Iron acquisition and storage by pathogenic fungi
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“Albomycin, a sideromycin produced by Streptomyces spp., protects mice from S. pneumoniae and Y. enterocolitica systemic infection (Pramanik et al., 2007).”
§ Iron in Infection and Immunity > V. Targeting microbial iron homeostasis for the treatment of infectious diseases
What the sources say
- Not checked yetcites Pramanik et al., 2007 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
In the paper
“By maintaining proper intracellular iron distribution, these compounds promote germination and resistance to oxidative stress, and are required for full virulence in a mouse model of invasive infection (Schrettl et al., 2007).”
§ Iron in Infection and Immunity > III. Iron acquisition and storage by pathogenic fungi
What the sources say
- Not checked yetcites Schrettl et al., 2007 source
Tests✓ Asserted: passes✓ Checkable: passes✓ Load-bearing: passes✓ Based: passesIt may state two separate claims as one.
In the paper
“Up to two-thirds of the world’s population are exposed to malaria annually, of which over 240 million will contract the disease and up to 2 million will die (Long et al., 2003).”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
What the sources say
- Not checked yetcites Long et al., 2003
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes✗ Load-bearing: fails✓ Based: passesProbably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Similarly, the synthetic siderophore-conjugated beta-lactam MC-1, is active against P. aeruginosa both in vitro and in vivo (McPherson et al., 2012).”
§ Iron in Infection and Immunity > V. Targeting microbial iron homeostasis for the treatment of infectious diseases
What the sources say
- Not checked yetcites McPherson et al., 2012 source
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The Plasmodium lifecycle includes an intra-erythrocytic stage, creating a unique opportunity to manipulate the richest reservoir of iron in the host.”
§ Iron in Infection and Immunity > IV. Iron homeostasis in human parasites
Strength:plain· not yet calibratedCausal:direct· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
- Rests on an assumption: Malaria is caused by one of five species of Plasmodium, with P. falciparum being the most virulent.
In the paper
“A. fumigatus, an important cause of invasive disease in immunocompromised patients, produces two intracellular siderophores: hyphal ferricrocin and conidial hydroxyferricrocin (Schrettl and Haas, 2011).”
§ Iron in Infection and Immunity > III. Iron acquisition and storage by pathogenic fungi
What the sources say
- Not checked yetcites Schrettl and Haas, 2011 source
Strength:plain· not yet calibratedTests✓ Asserted: passes✓ Checkable: passes? Load-bearing: unclear✓ Based: passesIt may state two separate claims as one.
The paper relies on this without support.
The vast majority of human pathogens require iron to sustain growth and colonize humans.
“The vast majority of human pathogens require iron to sustain growth and colonize humans.”
The paper relies on this without support.
Hookworms feed on human blood, using a hemolysin to lyse erythrocytes and anticoagulants to resist clotting.
“Hookworms feed on human blood, using a hemolysin to lyse erythrocytes and anticoagulants to resist clotting.”
The paper relies on this without support.
Malaria is caused by one of five species of Plasmodium, with P. falciparum being the most virulent.
“Malaria is caused by one of five species of Plasmodium, with P. falciparum being the most virulent.”
The paper relies on this without support.
Hemoglobin is ingested by intra-erythrocytic Plasmodia, transported to a food vacuole, and degraded to nutrient amino acids by a proteolytic cascade.
“Hemoglobin is ingested by intra-erythrocytic Plasmodia, transported to a specialized organelle known as the food vacuole, and degraded to nutrient amino acids by a proteolytic cascade.”
The paper relies on this without support.
Digestion of hemoglobin releases heme, which is toxic at elevated concentrations.
“However, digestion of hemoglobin also releases heme, which is toxic at elevated concentrations.”
The paper relies on this without support.
Plasmodia sequester heme into hemozoin, which accumulates in organs of infected individuals.
“To limit toxicity, Plasmodia sequester heme into a pigment known as hemozoin, which accumulates in the organs of infected individuals.”
The paper relies on this without support.
Non-Typhoid Salmonella (NTS) bacteremia is a common and often fatal complication of malaria.
“Non-Typhoid Salmonella (NTS) bacteremia is a common and often fatal complication of malaria.”
The paper relies on this without support.
Hemolysis during malaria induces the expression of HO-1, which protects host cells by detoxifying heme.
“Hemolysis during malaria induces the expression of HO-1, which protects host cells by detoxifying heme.”
The paper relies on this without support.
Lcn2 is abundantly produced during malaria.
“The siderophore-sequestering molecule Lcn2 is abundantly produced during malaria.”
The paper relies on this without support.
Siderophore conjugation decreases the minimum inhibitory concentration of several antibiotics, including beta-lactams, vancomycin, and fluoroquinolones.
“Siderophore conjugation decreases the minimum inhibitory concentration of several antibiotics, including beta-lactams, vancomycin, and fluoroquinolones.”
The paper relies on this without support.
The Na-APR-1 protease from human hookworm, Necator americanus, is essential for enzymatic activity to support blood feeding.
“The Na-APR-1 protease from human hookworm, Necator americanus, is essential for enzymatic activity to support blood feeding.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“Iron (Fe) is almost universally essential for life and is among the most fiercely fought over elements.118,119”
From Metals in Motion: Understanding Labile Metal Pools in Bacteria (2025) · cites this paper as [119]
Experimental — accuracy not yet measuredThe cited source says what the claim attributes to it.
In the citing paper
“Elevated concentrations of hepcidin increase iron sequestration by macrophages and decrease duodenal iron absorption during inflammatory reactions and thus reduces plasma iron levels (Cassat and Skaar, 2013; Noble, 2013).”
From Iron at the Centre of Candida albicans Interactions (2018) · cites this paper as Cassat and Skaar, 2013
The cited source says what the claim attributes to it.
In the citing paper
“In addition, iron can directly promote microbial growth (Cassat and Skaar, 2013), Zimmermann et al. (2010) reported an increase in enteric pathogens with iron supplementation.”
From Iron at the Centre of Candida albicans Interactions (2018) · cites this paper as Cassat and Skaar, 2013
The cited source says what the claim attributes to it.
In the citing paper
“The control over iron homeostasis is decisive in host–pathogen interaction (Schaible and Kaufmann, 2004; Nairz et al., 2010; Cassat and Skaar, 2013).”
From Iron at the interface of immunity and infection (2014) · cites this paper as Cassat and Skaar, 2013
The cited source says what the claim attributes to it.
In the citing paper
“Many microbes are highly dependent on a sufficient supply of iron and take up this metal by multiple and divergent pathways or steel it from iron deposition sites of the host (Winkelmann, 2002; Cassat and Skaar, 2013; Frawley et al., 2013).”
From Iron at the interface of immunity and infection (2014) · cites this paper as Cassat and Skaar, 2013
In the citing paper
“Rapid microbial multiplication and clonal selection allow microorganisms to evade conditions that restrict their growth, including iron sequestration by their hosts79.”
From Iron homeostasis in host defence and inflammation (2015) · cites this paper as [79]
In the citing paper
“Based on the decisive role of iron for both host immune system and microbe function, it is obvious that iron metabolism is significantly changed during the course of an infection (32, 167, 168).”
From Macrophage defense mechanisms against intracellular bacteria (2015) · cites this paper as [168]
In the citing paper
“These alterations of iron traffic are thought to result from defense strategy of the body to limit the availability of iron for invading pathogens (167–169), which lead to retention of iron in MΦs and an impaired iron absorption from the diet (170).”
From Macrophage defense mechanisms against intracellular bacteria (2015) · cites this paper as [168]
In the citing paper
“These mycobacterial siderophores can diffuse out of the phagloysome into the cytoplasm and are re-introduced by the bacteria to increase their access to iron, which is essential for the pathogenicity of M. tuberculosis (168, 206).”
From Macrophage defense mechanisms against intracellular bacteria (2015) · cites this paper as [168]
In the citing paper
“M. tuberculosis produces two classes of such siderophores, namely mycobactins and carboxymycobactins.”
From Macrophage defense mechanisms against intracellular bacteria (2015) · cites this paper as [168]
In the citing paper
“Invading pathogens thrive on free iron in the bloodstream to proliferate and advance their attack (Cassat and Skaar, 2013; Figure 3).”
From The Role of Iron Regulation in Immunometabolism and Immune-Related Disease (2019) · cites this paper as Cassat and Skaar, 2013
In the citing paper
“First, a range of intracellular microbes can resist killing by macrophages by gaining access to macrophage iron pools [30, 245].”
From “Pumping iron”—how macrophages handle iron at the systemic, microenvironmental, and cellular levels (2017) · cites this paper as [30]
In the citing paper
“Of note, deletion of genes regulating the expression of siderophores or other Fe uptake systems is often associated with reduced pathogen virulence, as illustrated for bacteria, for example, Staphylococci and Salmonella, or fungi, for example, Aspergillus, infections [6,7,32,33].”
From The Iron age of host–microbe interactions (2015) · cites this paper as [7]
In the citing paper
“Subsequent microbiology and molecular genetic studies established that nutritional immunity plays a pivotal role in defense against an array of pathogens, including bacteria, fungi, and parasites 3, 15.”
From Buried Treasure: Evolutionary Perspectives on Microbial Iron Piracy (2015) · cites this paper as [3]
In the citing paper
“Nutrient iron sequestration provides one such alternative mode of host defense against bacteria and eukaryotic pathogens [3].”
From Buried Treasure: Evolutionary Perspectives on Microbial Iron Piracy (2015) · cites this paper as [3]
In the citing paper
“Iron acquisition genes are now established as microbial virulence factors, in the sense that loss of these genes impairs pathogenicity without completely compromising organism viability 3, 86.”
From Buried Treasure: Evolutionary Perspectives on Microbial Iron Piracy (2015) · cites this paper as [3]
In the citing paper
“While the iron acquisition mechanisms of aerobic Gram-negative bacteria are known to be essential to their virulence (5), the host mechanisms of iron restriction in these infections are incompletely defined.”
From Hepcidin-mediated iron sequestration protects against bacterial dissemination during pneumonia (2017) · cites this paper as [5]
In the citing paper
“In response to infection, patients commonly develop hypoferremia (i.e., a decrease of iron in the circulation), a host response to limit iron availability to invading pathogens (3).”
From Iron- and Hepcidin-Independent Downregulation of the Iron Exporter Ferroportin in Macrophages during Salmonella Infection (2017) · cites this paper as [3]
In the citing paper
“Iron is an indispensable metal for the spread of virtually all human pathogens (3).”
From Iron- and Hepcidin-Independent Downregulation of the Iron Exporter Ferroportin in Macrophages during Salmonella Infection (2017) · cites this paper as [3]
In the citing paper
“where competition for limited Fe resources emerged as an evolutionary conserved host defense strategy against infection in plants (2) and animals (3–5) as well as a microbial virulence factor (6).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“Presumably, this explains why most pathogens evolved a variety of strategies aimed at diverting Fe from their hosts into their own metabolic pathways, while hosts co-evolved strategies to restrict pathogens from accessing Fe (3–6).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“This microbial Fe acquisition strategy is countered by reducing Fe concentration in plasma, a host defense strategy termed hypoferremia (3, 5).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“The cytosolic heme is used to extract Fe by heme oxygenases (3, 77).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“Gram-negative pathogenic bacteria such as Yersinia enterocolitica, Shigella, E. coli O157:H7, Yersinia pestis or Haemophilus influenzae, express outer membrane Hb and/or heme receptors, while Yersinia pestis, Pseudomonas aeruginosa and Serratia marcescens secret hemophores that bind extracellular heme (3, 77).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“Gram-negative pathogenic bacteria such as Yersinia enterocolitica, Shigella, E. coli O157:H7, Yersinia pestis or Haemophilus influenzae, express outer membrane Hb and/or heme receptors, while Yersinia pestis, Pseudomonas aeruginosa and Serratia marcescens secret hemophores that bind extracellular heme (3, 77).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“These are recognized by outer membrane TonB-dependent heme, hemoprotein and hemophore receptors, respectively, which shuttle heme to the bacterial periplasm to be transported into the cytosol by specific ABC protein-dependent periplasmic permeases (3, 77).”
From Innate Nutritional Immunity (2018) · cites this paper as [3]
In the citing paper
“Hb contains about two-thirds of the body's iron, and a single erythrocyte contains more than 280 million molecules of Hb [15,24].”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“bacterial pathogens have developed exquisite tactics to overcome iron limitation and elaborate high-affinity iron receptors and chelators”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“The limited access of pathogens to metals serves as an antimicrobial strategy and limits bacterial replication.”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“Heme makes up the greatest reservoir of iron in the host and serves as an iron source for many bacterial pathogens.”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“Its abundance and location in erythrocytes make Hb a rich heme source for pathogens.”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“The solubility of ferric iron in aerobic solution is exceedingly low”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“high-affinity iron-binding proteins, including transferrin, lactoferrin, albumin, and ferritin, sequester iron extracellularly and intracellularly”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“For instance, free iron rarely exists in the mammalian host.”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“Nutritional immunity has since been expanded to include the host processes that manipulate local levels of manganese, zinc, and other transition metals in order to metal starve or intoxicate the invading pathogens”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“Iron-binding proteins function to transport iron, protect host cells from iron-mediated oxidative damage, and keep iron from pathogens.”
From Heme Synthesis and Acquisition in Bacterial Pathogens (2016) · cites this paper as [15]
In the citing paper
“As with many pathogens (Schaible and Kaufmann, 2004), iron is essential for proliferation and virulence of S. aureus (Cassat and Skaar, 2013).”
From Staphylococcus aureus Targets the Duffy Antigen Receptor for Chemokines (DARC) to Lyse Erythrocytes (2015) · cites this paper as Cassat and Skaar, 2013
Citations
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Hepcidin Regulates Cellular Iron Efflux by Binding to Ferroportin and Inducing Its Internalization
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Metadata
- Authors
- Cassat, J. E., & Skaar, E. P.
- Affiliations
- Vanderbilt University Medical Center (US)
- Vanderbilt University (US)
- Corresponding author
- Skaar, Eric P.
- Venue
- Cell Host & Microbe
- Publisher
- Elsevier BV
- Published
- 2013-05
- Volume
- 13
- Issue
- 5
- Pages
- 509-519
- ISSN
- 1931-3128 (ISSN-L)
- PMC
- PMC3676888
- OpenAlex
- W2044678473
- Cited by
- 1,214
- References
- 97
- Citations by year
- 2026: 62 · 2025: 90 · 2024: 91 · 2023: 106 · 2022: 145 · 2021: 150 · 2020: 112 · 2019: 108 · 2018: 84 · 2017: 83 · 2016: 71 · 2015: 62 · 2014: 39 · 2013: 7
- FWCI
- 9.11
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Cell Host & Microbe (publishedVersion)
- PubMed (publishedVersion)
- Europe PMC (PubMed Central) (acceptedVersion)
- PubMed Central (acceptedVersion)
- Funders
- National Institute of Allergy and Infectious Diseases (L40 AI096516, R01AI069233, R01AI073843, R01AI101171)
- Topics
- Health Sciences › Medicine › Hematology › Iron Metabolism and Disorders
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Bacterial Genetics and Biotechnology
- Physical Sciences › Materials Science › Metals and Alloys › Iron and Steelmaking Processes
- Keywords
- iron metabolism, nutritional immunity, iron acquisition, iron regulation, innate immunity, host-pathogen interaction, iron redox chemistry, iron toxicity
- MeSH terms
- Animals, Bacteria, Communicable Diseases, Fungi, Homeostasis, Humans, Immunity, Innate, Iron, Models, Biological, Parasites
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