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Magnesium is an essential element in biological systems.

Magnesium
occurs typically as the Mg2+ ion. It is an essential mineral nutrient for life[1][2]
[3] and is present in every cell type in every organism. For example, ATP
(adenosine triphosphate), the main source of energy in cells, must be bound to
a magnesium ion in order to be biologically active. What is called ATP is often
actually Mg-ATP. [4]. Similarly, magnesium plays a role in the stability of all
polyphosphate compounds in the cells, including those associated with DNA
and RNA
Function
• A balance of magnesium is vital to the well being of all
organisms. Magnesium is a relatively abundant ion in the
lithosphere and is highly bioavailable in the hydrosphere.
This ready availability, in combination with a useful and
very unusual chemistry, may have led to its usefulness in
evolution as an ion for signalling, enzyme activation and
catalysis. However, the unusual nature of ionic
magnesium has also led to a major challenge in the use
of the ion in biological systems. Biological membranes
are impermeable to Mg2+ (and other ions) so transport
proteins must facilitate the flow of Mg2+, both into and
out of cells and intracellular compartments.
Biological range, distribution, and
regulation
• In animals it has been shown that different cell types maintain different concentrations of
magnesium.[5][6][7][8] It seems likely that the same is true for plants.[9][10] This suggests that
different cell types may regulate influx and efflux of magnesium in different ways based on their
unique metabolic needs. Interstitial and systemic concentrations of free magnesium must be
delicately maintained by the combined processes of buffering (binding of ions to proteins and
other molecules) and muffling (the transport of ions to storage or extracellular spaces[11]).
• In plants, and more recently in animals, magnesium has been recognized as an important
signaling ion, both activating and mediating many biochemical reactions. The best example of
this is perhaps the regulation of carbon fixation in chloroplasts in the Calvin cycle.[12][13]
• The importance of magnesium to proper cellular function cannot be overstated. Deficiency of the
nutrient results in disease in the affected organism. In single-celled organisms such as bacteria
and yeast, low levels of magnesium manifests in greatly reduced growth rates. In magnesium
transport knockout strains of bacteria, healthy rates are maintained only with exposure to very
high external concentrations of the ion.[14][15] In yeast, mitochondrial magnesium deficiency is
also leads to disease.[16]
• Plants deficient in magnesium show stress responses. The first observable signs of both
magnesium starvation and overexposure in plants is a decrease in the rate of photosynthesis.
This is due to the central position of the Mg++ ion in the chlorophyll molecule. The later effects of
magnesium deficiency on plants are a significant reduction in growth and reproductive viability.[3]
Magnesium can also be toxic to plants, although this is typically seen only in drought conditions.
[17][18]
Biological range, distribution, and
regulation
• Space-filling model of the chlorophyll a molecule, with the
Magnesium ion (bright green) visible at the center of the porphyrin
group
• In animals, magnesium deficiency (hypomagnesemia) is seen when
the environmental availability of magnesium is low. In ruminant
animals, particularly vulnerable to magnesium availability in pasture
grasses, the condition is known as ‘grass tetany’. Hypomagnesemia
is identified by a loss of balance due to muscle weakness.[19] A
number of genetically attributable hypomagnesemia disorders have
also been identified in humans.[20][21][22][23]
• Overexposure to magnesium may be toxic to individual cells, though
these effects have been difficult to show experimentally. In humans
the condition is termed hypermagnesemia, and is well documented,
though it is usually caused by loss of kidney function. In healthy
individuals, excess magnesium is rapidly excreted in the urine
Human health
• Magnesium deficiency in humans was first described in the medical literature in 1934.
The adult human daily nutritional requirement, which is affected by various factors
including gender, weight and size, is 300-400 mg/day. Inadequate magnesium intake
frequently causes muscle spasms, and has been associated with
cardiovascular disease, diabetes, high blood pressure, anxiety disorders, migraines,
osteoporosis and cerebral infarction[24]. Acute deficiency (see hypomagnesemia) is
rare, and is more common as a drug side effect (such as chronic alcohol or diuretic
use) than from low food intake per se, but it can also occur within people fed
intravenously for extended periods of time. The incidence of chronic deficiency
resulting in less than optimal health is debated.
• The DRI upper tolerated limit for supplemental magnesium is 350 mg/day (calculated
as milligrams (mg) of elemental magnesium in the salt). (Supplements based on
amino acid chelates, such as glycinate, lysinate etc., are much better tolerated by the
digestive system and do not have the side effects of the older compounds used, while
sustained release supplements prevent the occurrence of diarrhea.)[citation needed]
The most common symptom of excess oral magnesium intake is diarrhea. Since the
kidneys of adult humans excrete excess magnesium efficiently, oral magnesium
poisoning in adults with normal renal function is very rare. Infants, which have less
ability to excrete excess magnesium even when healthy, should not be given
magnesium supplements, except under a physician's care
Human health
Magnesium salts (usually in the form of magnesium sulfate or chloride
when given parenterally) are used therapeutically for a number of
medical conditions, see Epsom salts for a list of conditions which have
been treated with supplemental magnesium ion. Magnesium is
absorbed with reasonable efficiency (30% to 40%) by the body from
any soluble magnesium salt, such as the chloride or citrate.
Magnesium is similarly absorbed from Epsom salts, although the
sulfate in these salts adds to their laxative effect at higher doses.
Magnesium absorption from the insoluble oxide and hydroxide salts (
milk of magnesia) is erratic and of poorer efficiency, since it depends
on the neutralization and solution of the salt by the acid of the
stomach, which may not be (and usually is not) complete.
Magnesium orotate may be used as adjuvant therapy in patients on
optimal treatment for severe congestive heart failure, increasing
survival rate and improving clinical symptoms and patient's quality of
life
Nerve Conduction
• Magnesium can effect muscle relaxation
through direct action on the cell
membrane. Mg++ ions close certain types
of calcium channels, which conduct a
positively charged calcium ion into the
neuron. With an excess of magnesium,
more channels will be blocked and the
nerve will have less activity
Hypertension
• Magnesium-containing Epsom salts are especially used
in treating the hypertension of eclampsia. Even if the
case is not eclampsia, there may be antihypertensive
effects of having a substantial portion of the intake of
sodium chloride (NaCl) exchanged for e.g. magnesium
chloride; NaCl is an osmolite and increases
arginine vasopressin (AVP) release, which increases
extracellular volume and thus results in increased blood
pressure. However, not all osmolites have this effect on
AVP release[26], so with magnesium chloride, the
increase in osmolarity may not cause such a
hypertensive response
Food sources
• Some good sources of magnesium.
• Green vegetables such as spinach provide magnesium because of the abundance of chlorophyll molecules which
contain the ion. Nuts (especially cashews and almonds), seeds, and some whole grains are also good sources of
magnesium.
• Although many foods contain magnesium, it is usually found in low levels. As with most nutrients, daily needs for
magnesium are unlikely to be met by one serving of any single food. Eating a wide variety of fruits, vegetables,
and grains will help ensure adequate intake of magnesium.
• Because magnesium readily dissolves in water, refined foods, which are often processed or cooked in water and
dried, are generally poor sources of the nutrient. For example, whole-wheat bread has twice as much magnesium
as white bread because the magnesium-rich germ and bran are removed when white flour is processed. The
table of food sources of magnesium suggests many dietary sources of magnesium.
• "Hard" water can also provide magnesium, but "soft" water does not contain the ion. Dietary surveys do not
assess magnesium intake from water, which may lead to underestimating total magnesium intake and its
variability.
• Too much magnesium may make it difficult for the body to absorb calcium. Not enough magnesium can lead to
hypomagnesemia as described above, with irregular heartbeats, high blood pressure (a sign in humans but not
some experimental animals such as rodents), insomnia and muscle spasms (fasciculation). However, as noted,
symptoms of low magnesium from pure dietary deficiency are thought to be rarely encountered.
• Following are some foods and the amount of magnesium in them:
• spinach (1/2 cup) = 80 milligrams (mg)
• peanut butter (2 tablespoons) = 50 mg
• black-eyed peas (1/2 cup) = 45 mg
• milk: low fat (1 cup) = 40 mg
Biological chemistry

• Mg2+ is the fourth most abundant metal ion in cells (in
moles) and the most abundant free divalent cation — as
a result it is deeply and intrinsically woven into cellular
metabolism. Indeed, Mg2+-dependent enzymes appear
in virtually every metabolic pathway: specific binding of
Mg2+ to biological membranes is frequently observed,
Mg2+ is also used as a signalling molecule, and much of
nucleic acid biochemistry requires Mg2+, including all
reactions which require release of energy from ATP.[27]
[28][13] In nucleotides, the triple phosphate moiety of the
compound is invariably stabilized by association with
Mg2+ in all enzymic processes.
Chlorophyll
• In photosynthetic organisms Mg2+ has the
additional vital role of being the
coordinating ion in the chlorophyll
molecule. This role was discovered by R.
M. Willstätter, who received the Nobel
Prize in Chemistry 1915 for the purification
and structure of chlorophyll
Enzymes
• The chemistry of the Mg2+ ion, as applied to enzymes, uses the full
range of this ion’s unusual reaction chemistry to fulfill a range of
functions.[27][29][30][31] Mg2+ interacts with substrates, enzymes
and occasionally both (Mg2+ may form part of the active site). Mg2+
generally interacts with substrates through inner sphere
coordination, stabilising anions or reactive intermediates, also
including binding to ATP and activating the molecule to nucleophilic
attack. When interacting with enzymes and other proteins Mg2+
may bind using inner or outer sphere coordination, to either alter the
conformation of the enzyme or take part in the chemistry of the
catalytic reaction. In either case, because Mg2+ is only rarely fully
dehydrated during ligand binding, it may be a water molecule
associated with the Mg2+ that is important rather than the ion itself.
The Lewis acidity of Mg2+ (pKa 11.4) is used to allow both
hydrolysis and condensation reactions (most commonly phosphate
ester hydrolysis and phosphoryl transfer) that would otherwise
require pH values greatly removed from physiological values
Essential role in the biological
activity of ATP

• ATP (adenosine triphosphate), the main
source of energy in cells, must be bound
to a magnesium ion in order to be
biologically active. What is called ATP is
often actually Mg-ATP
Nucleic acids
• Nucleic acids have an important range of interactions with Mg2+.
The binding of Mg2+ to DNA and RNA stabilises structure; this can
be observed in the increased melting temperature (Tm) of double-
stranded DNA in the presence of Mg2+.[27] Additionally, ribosomes
contain large amounts of Mg2+ and the stabilisation provided is
essential to the complexation of this ribo-protein.[33] A large number
of enzymes involved in the biochemistry of nucleic acids bind Mg2+
for activity, using the ion for both activation and catalysis. Finally,
the autocatalysis of many ribozymes (enzymes containing only
RNA) is Mg2+ dependent (e.g. the yeast mitochondrial group II self
splicing introns[34]).
• Magnesium ions can be critical in maintaining the positional integrity
of closely clustered phosphate groups. These clusters appear in
numerous and distinct parts of the cell nucleus and cytoplasm. For
instance hexahydrated Mg2+ ions bind in the deep major groove
and at the outer mouth of A-form nucleic acid duplexes
Cell membranes and walls
• Biological cell membranes and cell walls are polyanionic surfaces.
This has important implications for the transport of ions, particularly
because it has been shown that different membranes preferentially
bind different ions.[27] Both Mg2+ and Ca2+ regularly stabilise
membranes by the cross-linking of carboxylated and phosphorylated
head groups of lipids. However, the envelope membrane of E. coli
has also been shown to bind Na+, K+, Mn2+ and Fe3+. The
transport of ions is dependent on both the concentration gradient of
the ion and the electric potential (ΔΨ) across the membrane, which
will be affected by the charge on the membrane surface. For
example, the specific binding of Mg2+ to the chloroplast envelope
has been implicated in a loss of photosynthetic efficiency by the
blockage of K+ uptake and the subsequent acidification of the
chloroplast stroma.[12]
Proteins
• The Mg2+ ion tends to bind only weakly to proteins (Ka ≤ 105[27])
and this can be exploited by the cell to switch enzymatic activity on
and off by changes in the local concentration of Mg2+. Although the
concentration of free cytoplasmic Mg2+ is on the order of 1 mmol/L,
the total Mg2+ content of animal cells is 30 mmol/L[36] and in plants
the content of leaf endodermal cells has been measured at values
as high as 100 mmol/L (Stelzer et al., 1990), much of which is
buffered in storage compartments. The cytoplasmic concentration of
free Mg2+ is buffered by binding to chelators (e.g. ATP), but also
more importantly by storage of Mg2+ in intracellular compartments.
The transport of Mg2+ between intracellular compartments may be
a major part of regulating enzyme activity. The interaction of Mg2+
with proteins must also be considered for the transport of the ion
across biological membranes
Importance in drug binding
• An article[37] investigating the structural basis of
interactions between clinically reelevant
antibiotics and the 50S ribosome appeared in
Nature in October 2001. High resolution x-ray
crystallography established that these antibiotics
only associate with the 23S rRNA of a ribosomal
subunit, and no interactions are formed with a
subunit's protein portion. The article stresses
that the results show "the importance of putative
Mg2+ ions for the binding of some drugs
Plant physiology of magnesium

• The previous sections have dealt in detail
with the chemical and biochemical aspects
of Mg2+ and its transport across cellular
membranes. This section will apply this
knowledge to aspects of whole plant
physiology, in an attempt to show how
these processes interact with the larger
and more complex environment of the
multicellular organism
Nutritional requirements and
interactions
• Mg2+ is essential for plant growth and is present in higher plants in amounts on the order of 80 μmol g-1 dry
weight.[3] The amounts of Mg2+ vary in different parts of the plant and are dependent upon nutritional status. In
times of plenty, excess Mg2+ may be stored in vascular cells (Stelzer et al., 1990;[10] and in times of starvation
Mg2+ is redistributed, in many plants, from older to newer leaves.[3][51]
• Mg2+ is taken up into plants via the roots. Interactions with other cations in the rhizosphere can have a significant
effect on the uptake of the ion.(Kurvits and Kirkby, 1980;[52] The structure of root cell walls is highly permeable to
water and ions, and hence ion uptake into root cells, can occur anywhere from the root hairs to cells located
almost in the centre of the root (limited only by the Casparian strip). Plant cell walls and membranes carry a great
number of negative charges and the interactions of cations with these charges is key to the uptake of cations by
root cells allowing a local concentrating effect.[53] Mg2+ binds relatively weakly to these charges, and can be
displaced by other cations, impeding uptake and causing deficiency in the plant.
• Within individual plant cells the Mg2+ requirements are largely the same as for all cellular life; Mg2+ is used to
stabilise membranes, is vital to the utilisation of ATP, is extensively involved in the nucleic acid biochemistry, and
is a cofactor for many enzymes (including the ribosome). Also, Mg2+ is the coordinating ion in the chlorophyll
molecule. It is the intracellular compartmentalisation of Mg2+ in plant cells that leads to additional complexity.
Four compartments within the plant cell have reported interactions with Mg2+. Initially Mg2+ will enter the cell into
the cytoplasm (by an as yet unidentified system), but free Mg2+ concentrations in this compartment are tightly
regulated at relatively low levels (≈2 mmol/L) and so any excess Mg2+ is either quickly exported or stored in the
second intracellular compartment, the vacuole.[54] The requirement for Mg2+ in mitochondria has been
demonstrated in yeast[55] and it seems highly likely that the same will apply in plants. The chloroplasts also
require significant amounts of internal Mg2+, and low concentrations of cytoplasmic Mg2+.[56][57] In addition, it
seems likely that the other subcellular organelles (e.g. Golgi, endoplasmic reticulum, etc) also require Mg2
Magnesium, chloroplasts and
photosynthesis
• Mg2+ is the coordinating metal ion in the chlorophyll molecule, and in plants
where the ion is in high supply about 6% of the total Mg2+ is bound to
chlorophyll.[3][61][62] Thylakoid stacking is stabilised by Mg2+ and is
important for the efficiency of photosynthesis, allowing phase transitions to
occur.[63]
• Mg2+ is probably taken up into chloroplasts to the greatest extent during the
light induced development from proplastid to chloroplast or etioplast to
chloroplast. At these times the synthesis of chlorophyll and the biogenesis
of the thylakoid membrane stacks absolutely require the divalent cation.[64]
[65]
• Whether Mg2+ is able to move into and out of chloroplasts after this initial
developmental phase has been the subject of several conflicting reports.
Deshaies et al. (1984) found that Mg2+ did move in and out of isolated
chloroplasts from young pea plants,[66] but Gupta and Berkowitz (1989)
were unable to reproduce the result using older spinach chloroplasts.[67]
Deshaies et al. had stated in their paper that older pea chloroplasts showed
less significant changes in Mg2+ content than those used to form their
conclusions. Perhaps the relative proportion of immature chloroplasts
present in the preparations might explain these observations.
Magnesium, chloroplasts and
photosynthesis
• The metabolic state of the chloroplast changes considerably between night and day.
During the day the chloroplast is actively harvesting the energy of light and converting
it into chemical energy. The activation of the metabolic pathways involved comes
from the changes in the chemical nature of the stroma on the addition of light. H+ is
pumped out of the stroma (into both the cytoplasm and the lumen) leading to an
alkaline pH.[68][69] Mg2+ (along with K+) is released from the lumen into the stroma,
in an electroneutralisation process to balance the flow of H+.[70][71][72][73] Finally,
thiol groups on enzymes are reduced by a change in the redox state of the stroma.
[74] Examples of enzymes activated in response to these changes are fructose 1,6-
bisphosphatase, sedoheptulose bisphosphatase and ribulose-1,5-bisphosphate
carboxylase.[3][30][74] During the dark period, if these enzymes were active a
wasteful cycling of products and substrates would occur.
• Two major classes of the enzymes that interact with Mg2+ in the stroma during the
light phase can be identified.[30] Firstly, enzymes in the glycolytic pathway most often
interact with two atoms of Mg2+. The first atom is as an allosteric modulator of the
enzymes’ activity, while the second forms part of the active site and is directly
involved in the catalytic reaction. The second class of enzymes include those where
the Mg2+ is complexed to nucleotide di- and tri-phosphates (ADP and ATP) and the
chemical change involves phosphoryl transfer. Mg2+ may also serve in a structural
maintenance role in these enzymes (e.g. enolase).
Magnesium is also used:
• To remove sulfur from iron and steel.
• To refine titanium in the Kroll process.
• To photoengrave plates in the printing industry.
• To combine in alloys, where this metal is essential for airplane and missile
construction.
• In the form of turnings or ribbons, to prepare Grignard reagents, which are
useful in organic synthesis.
• As an alloying agent, improving the mechanical, fabrication and welding
characteristics of aluminium.
• As an additive agent in conventional propellants and the production of
nodular graphite in cast iron.
• As a reducing agent for the production of uranium and other metals from
their salts.
• As a desiccant, since it easily reacts with water.
• As a sacrificial (galvanic) anode to protect underground tanks, pipelines,
buried structures, and water heaters.
magnesium compounds

• The magnesium ion is necessary for all life (see magnesium in biology), so
magnesium salts are an additive for foods, fertilizers (Mg is a component of
chlorophyll), and culture media.
• Magnesium hydroxide is used in milk of magnesia, its chloride, oxide,
gluconate, malate, orotate and citrate used as oral magnesium
supplements, and its sulfate (Epsom salts) for various purposes in medicine,
and elsewhere (see the article for more). Oral magnesium supplements
have been claimed to be therapeutic for some individuals who suffer from
Restless Leg Syndrome (RLS).[citation needed]
• Magnesium borate, magnesium salicylate and magnesium sulfate are used
as antiseptics.
• Magnesium bromide is used as a mild sedative (this action is due to the
bromide, not the magnesium).
• Dead-burned magnesite is used for refractory purposes such as brick and
liners in furnaces and converters.
• Magnesium carbonate (MgCO3) powder is also used by athletes, such as
gymnasts and weightlifters, to improve the grip on objects – the apparatus
or lifting bar
magnesium compounds
• Magnesium stearate is a slightly flammable white powder
with lubricative properties. In pharmaceutical technology
it is used in the manufacturing of tablets, to prevent the
tablets from sticking to the equipment during the tablet
compression process (i.e., when the tablet's substance is
pressed into tablet form).
• Magnesium sulfite is used in the manufacture of paper (
sulfite process).
• Magnesium phosphate is used to fireproof wood for
construction.
• Magnesium hexafluorosilicate is used in mothproofing of
textiles
Biological role
• Due to the important interaction between phosphate and
magnesium ions, magnesium ions are essential to the basic
nucleic acid chemistry of life, and thus are essential to all cells of all
known living organisms. Over 300 enzymes require the presence of
magnesium ions for their catalytic action, including all enzymes
utilizing or synthesizing ATP, or those which use other nucleotides
to synthesize DNA and RNA. ATP exists in cells normally as a
chelate of ATP and a magnesium ion.
• Plants have an additional use for magnesium in that chlorophylls are
magnesium-centered porphyrins. Magnesium deficiency in plants
causes late-season yellowing between leaf veins, especially in older
leaves, and can be corrected by applying Epsom salts (which is
rapidly leached), or else crushed dolomitic limestone to the soil.
Food sources of magnesium
• (Magnesium is a vital component of a healthy human diet.
Human magnesium deficiency including conditions which show few overt symptoms)
is relatively common, with only 32% of the United States meeting the RDA-DRI,[9]
and has been implicated in the development of a number of human illnesses such as
asthma, osteoporosis, and ADHD.[10]
• Adult human bodies contain about 24 grams of magnesium, with 60% in the skeleton,
39% intracellular (20% in skeletal muscle), and 1% extracellular. Serum levels are
typically 0.7 – 1.0 mmol/L. Serum magnesium levels may appear normal even in
cases of underlying intracellular deficiency, although no known mechanism maintains
a homeostatic level in the blood other than renal excretion of high blood levels.
Intracellular magnesium is correlated with intracellular potassium. Magnesium is
absorbed in the gastrointestinal tract, with more absorbed when status is lower. In
humans, magnesium appears to facilitate calcium absorption. Low and high protein
intake inhibit magnesium absorption, and other factors such as phosphate, phytate,
and fat affect absorption. Absorbed dietary magnesium is largely excreted through
the urine, although most magnesium "administered orally" is excreted through the
feces.[11] Magnesium status may be assessed roughly through serum and
erythrocyte Mg concentrations and urinary and fecal excretion, but intravenous
magnesium loading tests are likely the most accurate and practical in most people.
[12] In these tests, magnesium is injected intravenously; a retention of 20% or more
indicates deficiency.[13] Other nutrient deficiencies are identified through biomarkers,
but none are established for magnesium.[14]
Food sources of magnesium
• Spices, nuts, cereals, coffee, cocoa, tea,
and vegetables (especially green leafy
ones) are rich sources of magnesium.
Observations of reduced dietary
magnesium intake in modern Western
countries as compared to earlier
generations may be related to food
refining and modern fertilizers which
contain no magnesium