UPTAKE

FORMS,ASSIMILATION,ME
TABOLISM AND ROLE OF
SULFUR IN PLANTS
 Sulfur Assimilation
Sulfur is an essential element for growth and physiological functioning of plants
,however, its content strongly varies between plant species and it ranges from 0.1 to
6 % of the plants' dry weight. Sulfate taken up by the roots is the major sulfur source
for growth, though it has to be reduced to sulfide before it is further metabolized.
Root plastids contain all sulfate reduction enzymes, however, the reduction of
sulfate to sulfide and its subsequent incorporation into cysteine takes predominantly
place in the shoot in the chloroplast, Cysteine is the precursor or reduced sulfur
donor of most other organic sulfur compounds in plants. The predominant
proportion of the organic sulfur is present in the protein fraction (up to 70 % of total
sulfur), as cysteine and methionine residues. Cysteine and methionine are highly
significant in the structure, conformation and function of proteins. Plants contain a
large variety of other organic sulfur compounds, as thiols (glutathione), sulfolipids
and secondary sulfur compounds (alliins, glucosinolates, phytochelatins), which play
an important role in physiology and protection against environmental stress and
pests. Sulfur compounds are also of great importance for food quality and for the
production of phyto-pharmaceutics. Sulfur deficiency will result in the loss of plant
production, fitness and resistance to environmental stress and pests.
Sulfate reduction and assimilation in plants

APS, adenosine 5'phosphosulfate;

Fdred, Fdox, reduced and oxidized
ferredoxin; RSH, RSSR, reduced
and oxidized glutathione
 Sulfate uptake by plants
Sulfate is taken up by the roots with high affinity and the maximal sulfate uptake rate is
generally already reached at sulfate levels of 0.1 mM and lower. The uptake of sulfate by the
roots and its transport to the shoot is strictly controlled and it appears to be one of the primary
regulatory sites of sulfur assimilation. Sulfate is actively taken up across the plasma membrane
of the root cells, subsequently loaded into the xylem vessels and transported to the shoot by the
transpiration stream. The uptake and transport of sulfate is energy dependent (driven by a
proton gradient generated by ATPases) through a proton/sulfate co-transport. In the shoot the
sulfate is unloaded and transported to the chloroplasts where it is reduced. The remaining
sulfate in plant tissue is predominantly present in the vacuole, since the concentration of sulfate
in the cytoplasm is kept rather constant. Distinct sulfate transporter proteins mediate the
uptake, transport and subcellular distribution of sulfate. According to their cellular and
subcellular gen expression, and possible functioning the sulfate transporters gene family has
been classified in up to 5 different groups. Some groups are expressed exclusively in the roots
or shoots or expressed both in the roots and shoots. Group 1 are 'high affinity sulfate
transporters', which are involved in the uptake of sulfate by the roots. Group 2 are vascular
transporters and are 'low affinity sulfate transporters'. Group 3 is the so-called 'leaf group',
however, still little is known about the characteristics of this group. Group 4 transporters are
involved in the efflux of sulfate from the vacuoles, whereas the function of Group 5 sulfate
transporters is not known yet. Regulation and expression of the majority of sulfate transporters
are controlled by the sulfur nutritional status of the plants. Upon sulfate deprivation, the rapid
decrease in root sulfate is regularly accompanied by a strongly enhanced expression of most
sulfate transporter genes (up to 100-fold), accompanied by a substantially enhanced sulfate
uptake capacity. It is not yet solved, whether sulfate itself or metabolic products of the sulfur
assimilation (O-acetylserine, cysteine, glutathione) act as signals in the regulation of sulfate
uptake by the root and its transport to the shoot, and in the expression of the sulfate
transporters involved.
Forms of sulfur uptake
 sulfur compounds in plants
Cysteine
Sulfide is incorporated into cysteine, catalyzed by O-acetyl serine (thiol)
lyase, with O-acetyl serine as substrate. The synthesis of O-acetyl serine is
catalyzed by serine acetyl transferase and together with O-acetyl serine
(thiol) lyase it is associated as enzyme complex named cysteine synthase.
The formation of cysteine is the direct coupling step between sulfur (sulphur
metabolism) and nitrogen assimilation in plants. Cysteine is sulfur donor for
the synthesis of methionine, the major other sulfur-containing amino acid
present in plants. This happens through the transsulfuration pathway and
the methylation of homocysteine. Both cysteine and methionine are sulfur-
containing amino acids and are of great significance in the structure,
conformation and function of proteins and enzymes, but high levels of these
amino acids may also be present in seed storage proteins. The thiol groups
of the cysteine residues in proteins can be oxidized resulting in disulfide
bridges with other cysteine side chains (and form cystine) and/or linkage of
polypeptides. Disulfide bridges (disulfide bonds) make an important
contribution to the structure of proteins. The thiol groups are also of great
importance in substrate binding of enzymes, in metal-sulfur clusters in
proteins (e.g. ferredoxins) and in regulatory proteins (e.g. thioredoxins).
 sulfur compounds in plants
Glutathione
Glutathione or its homologues, e.g. homoglutathione in Fabaceae;
hydroxymethylglutathione in Poaceae are the major water-soluble non-protein thiol
compounds present in plant tissue and account for 1-2 % of the total sulfur. The content
of glutathione in plant tissue ranges from 0.1 - 3 mM. Cysteine is the direct precursor for
the synthesis of glutathione (and its homologues). First, γ- glutamyl cysteine is
synthesized from cysteine and glutamate catalyzed by gamma-glutamyl cysteine
synthetase. Second, glutathione is synthesized from γ-glutamyl cysteine and glycine (in
glutathione homologues, β- alanine or serine) catalyzed by glutathione synthetase. Both
steps of the synthesis of glutathione are ATP dependent reactions. Glutathione is
maintained in the reduced form by an NADPH-dependent glutathione reductase and the
ratio of reduced glutathione (GSH) to oxidized glutathione (GSSG) generally exceeds a
value of 7. Glutathione fulfils various roles in plant functioning. In sulfur metabolism it
functions as reductant in the reduction of APS to sulfite. It is also the major transport
form of reduced sulfur in plants. Roots likely largely depend for their reduced sulfur
supply on shoot/root transfer of glutathione via the phloem, since the reduction of
sulfur occurs predominantly in the chloroplast. Glutathione is directly involved in the
reduction and assimilation of selenite into selenocysteine. Furthermore, glutathione is
of great significance in the protection of plants against oxidative and environmental
stress and it depresses/scavenges the formation of toxic reactive oxygen species, e.g.
superoxide, hydrogen peroxide and lipid hydroperoxides .
 Sulfur Deficiency Symptoms
Sulfur deficiency symptoms in most crops first appear on younger, upper
leaves. Signs unique to canola are delayed and prolonged flowering, pale,
whitish flowers; erect leaves on the upper parts of the plant; young leaves cup
inward and develop a pink, reddish or even purple color on the underside; seed
pods will be small and poorly filled.

SOIL TESTING. This will provide a general indication of a soil's sulfur status,
but it's best if combined with an early-season plant analysis (tissue test).
Because sulfur is a mobile soil nutrient, Canadian specialists recommend
taking soil samples from increments of 0-6 inches, 6-12 inches, and 12-24
inches. North Dakota specialist Dave Franzen suggests a 0-6 inch and 6-24 inch
sampling. On hilly acreage, it's a good idea to take separate samples from mid-
slopes and low-lying areas because sulfur deficiency is more common on
knolls.

"The degree of sulfur deficiency can vary throughout the field," says Jack
Dobb, agronomist with the Ministry of Agriculture and Fisheries, British
Columbia. "Even if the average reading indicates that a soil contains enough
sulfur, the crop may still benefit from sulfur fertilization."
BIOGEOCHEMICAL CYCLE OF SULFUR
THE SULFUR CYCLE
THE GLOBAL SULFUR CYCLE