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Uptake and transport of calcium in plants

Article  in  Zhi wu sheng li yu fen zi sheng wu xue xue bao = Journal of plant physiology and molecular biology · July 2005
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　Ｊｏｕｒｎａｌ　 ｏ ｆ　 Ｐ ｌａｎｔ　 Ｐ ｈｙｓｉｏｌｏｇｙ　 ａ ｎｄ　 Ｍ ｏｌｅｃｕｌａｒ　 Ｂ ｉｏｌｏｇｙ　　 ２ ００５， 　 ３ １ 　 （ ３）： 　 ２ ２７ －２３４
综　述 Review
Uptake and Transport of Calcium in Plants
YANG Hong-Qiang*, JIE Yu-Ling
(College of Horticulture, Shandong Agricultural University, Tai’an 271018, China)
Abstract: Recently, research on Ca 2+ transport in
plants has been focused on cellular and molecular level.
But the uptake, transport and distribution are also very
important for calcium to accomplish its function at
whole plant level. There are many cells along the way
of transport of Ca2+ from root to shoot, and Ca2+ passes
either through the cytoplasm of cells linked by plas-
modesmata (the symplast) or through the spaces be-
tween cells (the apoplast), which include Ca2+ uptake
by root cells, Ca2+ transport from root cortex to and
through the xylem, and then out of it into leaves or
fruits. Ca2+ channels, Ca2+/H+ antiporter and Ca2+-AT-
Pase play roles in the uptake and transport of Ca2+ in
root cells. To be transported from root surface to xylem,
Ca2+ needs to traverse endodermal cells and xylem
parenchyma cells. Endodermal Casparian band, the
main barrier for the apoplastic movement of ions into
the stele, compels some Ca2+ to enter root symplast
through Ca2+ channels in endodermal cells and then
reach xylem parenchyma. Ca2+-ATPase may drive Ca2+
into the stelar apoplast from xylem parenchyma. Some
Ca2+ effuses from endodermal cell and then get to xy-
lem through apoplastic pathway. Ca2+ is transported
in plant xylem vessel in chelate form and the speed of
water flow is the key factor Ca2+ transport via xylem
in trunk. There are both apoplastic and symplastic path-
ways of Ca2+ transport in fruit or leaf tissue too.
Key words: calcium; root; uptake and transport; Ca2+-ATPase;
xylem
Plant requires calcium in large amounts; the cal-
cium content of healthy plant tissues is generally more
than 0.1%–1% of dry matter (White 2001). Calcium
plays a variety of structural roles. Ca-deficiency disor-
ders often occur in horticulture, such as “tipburn” or
227
“brown heart” of leafy vegetables, “black heart” of
celery, “blossom end rot” of watermelon, pepper and
tomato fruits, “bitter pit” of apple and “empty pod” of
peanut, though calcium deficiency is rare in nature.
Other physiological disorders, such as “cracking” in
tomato, cherry and apple fruit following high humid-
ity or rainfall, often occur in tissues without enough
calcium (Shear, 1975). These disorders are mostly
caused by slow absorption and poor distribution of Ca2+
after transport. Calcium functions as a second mes-
senger in plant growth, development and adaptation
to environment (White and Broadley 2003).
Recently, research on Ca2+ in plants has been fo-
cused on the cellular and molecular levels. But the
uptake, transport and distribution are also very impor-
tant for calcium accomplishing its function at whole
plant level. The movement of Ca2+ is more difficult
than other nutrient elements; the rate of transport is
more likely to limit the supply. This article provides
an overview of recent works on Ca2+ uptake and deliv-
ery from root to leaf or fruit through apoplastic and
symplastic pathways.
1 Calcium uptake in root cell
The epidermal cells and their elongated projec-
tion (root hairs) contact with the soil solution, Ca2 +
can enter directly to them through channels. Root hairs
are specialized for the uptake of nutrients from the soil
by forming a very large surface area for absorption
per unit root length. The major role of root hair is in
the uptake of poorly mobile nutrients. It has been found
Ｒｅｃｅｉｖｅｄ　２００４－０８－０２，　Ａｃｃｅｐｔｅｄ　２００５－０３－０２．
This work was supported by the National Natural Science Foundation
of China (Nos. 30170655, 30270923).
* E-mail: labft@sdau.edu.cn; yhqsd@yahoo.com.cn
228 　植物生理与分子生物学学报
by using micro-electrodes that Ca2+ influx is highest
at the tips of root hairs (Gilroy and Jones 2000).
Most nutrients need to move through the sym-
plast from epidermal cells to the cortex and then to the
stele. Casparian strip of the endodermis is a major bar-
rier for the apoplastic movement and Ca2+ must enter
the cytoplasm of the endodermal cell through the Ca2+
channels in plasmolemma. Many kinds of Ca2+ chan-
nels have been found in root cells, such as depolariza-
tion-activated Ca2+ channels (White 1998, 2000), hy-
perpolarization-activated Ca2+ channels (Kiegle et al.
2000; Véry and Davies 2000) and voltage-insensitive
cation (VIC) channels (Davenport and Tester 2000)
There are also outward-rectifying cation (KORC or
NORC) channels (de Boer 1999), mechanosensitive
(stretch-activated) and second messenger-activated
Ca2+ channels (White 1998; Leng et al. 1999). Ion chan-
nels play roles in movement of mineral nutrients. The
hyperpolarization-activated Ca2+ channels have been
found only in cells of the elongation zone of the grow-
ing root apex of Arabidopsis, a region with a high de-
mand for Ca 2+ uptake for cell division and cell
e xpans ion. A la rge r C a 2+ flow a c tiva te d by
hyperpolarisation has also been found in cells from
the endodermis (Kiegle et al. 2000).
For the uptake and transfer of nutrients to the
xylem in roots, not only Ca2+ channels in plasmolemma,
but also tonoplast plays an important role. There are
Ca2+/H+ antiporter and Ca2+-ATPase in tonoplast, which
can transport Ca2+ to the vacuole for regulating ion
concentrations in the cytosol and hence in the symplast.
Plant vacuoles are unique among eukaryotic organelles
in having two kinds of proton pumps, one driven by
H+-ATPase (V-ATPase) and the other by H+-pyrophos-
phatase (V-PPase). There numerous proton pumps are
abundant in the tonoplast. The proton gradient estab-
lished by the tonoplast H+-pumps are used to drive the
H+-coupled transport systems. One of these is the Ca2+/
H+ antiporter (Paul and Russel 2000), which has a low
affinity with Ca2+ and a high capacity for Ca2+ pumping,
and removes excess Ca2+ from the cytosol to lower the
Ca2+ concentration to micromolar levels. The other is
Ca2+-ATPase, which has high affinity with Ca2+ and
acts to lower the cytosolic Ca2+ level further. Ca2+ -
　３１ 卷
ATPase and Ca2+/H+ antiporter coexist in the vacuolar
membrane (Ueoka-Nakanishi et al. 1999).
2 Transport from root to trunk xylem
2.1 Root cell types and endodermis function
The root is composed of many cell types, of which
each is specialized in a particular task (Maathuis et al.
1998). The outer cell layers of the root, the epidermis
and cortex, are involved primarily in the acquisition of
water and mineral nutrients. These cells are respon-
sible for the loading, unloading and long-distance trans-
port of solutes in the xylem and phloem, through the
endodermal cell layer. The part of the root outside the
plasma membrane forms the apoplast of root. This in-
cludes cell walls, intercellular spaces and the lumena
of tracheary elements. The interconnected cytoplasm
of root cells forms the symplast. Solutes may reach
xylem either symplastically, by entering root cells and
moving from cell to cell through plasmodesmata, or
apoplas tic ally, without tra ve rsing any plas ma
membrane. In mature root, a separation of cortical and
stelar apoplasts is formed by the presence of the
Casparian band. This structure, which is present on the
anticlinal (transverse and radial longitudinal) walls of
endodermal cells, is present in all roots of vascular
plants and act as a barrier between the parts of apoplast
inside and outside the endodermis.
The development of the endodermis can be di-
vided into three stages (Fig.1). At the first one, a
Casparian band shapes up within the anticlinal walls
of the endodermal cells (State I), to which plasma mem-
brane of these cells becomes firmly attached. The
Casparian band is composed not only of lignin, but
also some suberin, which blocks apoplastic (cell wall)
movement of Ca2+ to xylem. So any solute in the pe-
ripheral region can pass into the inner region only
through the protoplasm of the cells of the Casparian
band. When endodermis develops to State II and III,
layers of aliphatic suberin polymers are deposited over
the inner surface of the walls surrounding endodermal
cells and the layers of suberin form a lamellar structure,
which prevents endodermal cells from taking up sol-
utes from the apoplast (Peterson and Cholewa 1998).
During lateral root formation, the endodermal cells
divide and the newly formed radial walls where
３期 杨洪强等：　植物钙素吸收和运转（英文） 229

Ｆ ｉ ｇ ． １ 　 　 　　Ｓｃｈｅｍａｔｉｃ　ｒｏｕｔｅｓ　ｏｆ　ｃａｌｃｉｕｍ　ｔｒａｖｅｒｓｅ　ｆｒｏｍ　ｃｏｒｔｅｘ　ｔｏ　ｘｙｌｅｍ　ｉｎ　ｒｏｏｔ
Ｔｈｅ　ａｐｉｃａｌ　ｚｏｎｅ　ｏｆ　ｒｏｏｔ　ｉｓ　ｔｈｅ　ｍａｉｎ　ｐａｒｔ　ｏｆ　ｃａｌｃｉｕｍ　ａｂｓｏｒｐｔｉｏｎ．　Ｆｏｒ　ｃａｌｃｉｕｍ　ｍｏｖｅ　ｆｒｏｍ　ｒｏｏｔ　ｓｕｒｆａｃｅ　ｔｏ　ｘｙｌｅｍ，　ｉｔ　ｎｅｅｄｓ　ｔｏ　ｔｒａｖｅｒｓｅ
ｅｎｄｏｄｅｒｍａｌ　ｃｅｌｌｓ　ａｎｄ　ｘｙｌｅｍ　ｐａｒｅｎｃｈｙｍａ　ｃｅｌｌｓ．　Ｔｈｅｒｅ　ａｒｅ　ｔｈｒｅｅ　ｓｔａｔｕｓ　ｔｈａｔ　ｉｍｐａｃｔ　ｏｎ　ｔｈｅ　ｐａｔｈｗａｙ　ｏｆ　ｃａｌｃｉｕｍ　ｍｏｖｅｍｅｎｔ　ｔｏ　ｔｈｅ　ｘｙｌｅｍ
ａｃｃｏｒｄｉｎｇ　ｔｏ　Ｃａｓｐａｒｉａｎ　ｂａｎｄ　ｄｅｖｅｌｏｐｍｅｎｔ　（Ｗｈｉｔｅ　２００１）．　Ｉｎ　Ｓｔａｔｅ　Ｉ，　ｔｈｅ　Ｃａｓｐａｒｉａｎ　ｂａｎｄ　ｄｅｖｅｌｏｐｓ　ａｓ　ａ　ｍｏｄｉｆｉｃａｔｉｏｎ　ｏｆ　ｔｈｅ　ａｎｔｉｃｌｉｎａｌ
ｗａｌｌｓ　ｂｅｔｗｅｅｎ　ｅｎｄｏｄｅｒｍａｌ　ｃｅｌｌｓ．　Ｉｎ　ｔｈｉｓ　ｓｔａｔｅ，　ｃａｌｃｉｕｍ　ｍｏｖｅｍｅｎｔ　ｔｏ　ｔｈｅ　ｘｙｌｅｍ　ｉｓ　ｌｉｍｉｔｅｄ　ｉｎ　ｔｈｅ　ａｐｏｐｌａｓｔ　（ｃｅｌｌ　ｗａｌｌ），　ａｎｄ　ｆｏｒｃｅｄ　ｔｏ　ｅｎｔｅｒ
ｔｈｅ　ｒｏｏｔ　ｓｙｍｐｌａｓｔ　ｔｈｒｏｕｇｈ　Ｃａ２＋　ｃｈａｎｎｅｌｓ　ａｎｄ　ｐｕｍｐｅｄ　ｉｎｔｏ　ｔｈｅ　ｘｙｌｅｍ　ａｐｏｐｌａｓｔ　ｂｙ　Ｃａ２＋－ＡＴＰａｓｅ．　Ｉｎ　Ｓｔａｔｅ　ＩＩ，　ｌａｙｅｒｓ　ｏｆ　ｓｕｂｅｒｉｎ　ｐｏｌｙｍｅｒｓ
ａｒｅ　ｄｅｐｏｓｉｔｅｄ　ｏｖｅｒ　ｔｈｅ　ｅｎｔｉｒｅ　ｉｎｎｅｒ　ｓｕｒｆａｃｅ　ｏｆ　ｔｈｅ　ｗａｌｌｓ　ｓｕｒｒｏｕｎｄｉｎｇ　ｅｎｄｏｄｅｒｍａｌ　ｃｅｌｌｓ　ａｎｄ　ｌａｔｅｒａｌ　ｒｏｏｔｓ　ｏｆｔｅｎ　ｅｍｅｒｇｅ．　Ｉｎ　Ｓｔａｔｅ　ＩＩＩ，　ｔｈｅ
ｅｎｄｏｄｅｒｍａｌ　ｃｅｌｌ　ｉｓ　ｂｕｒｉｅｄ　ｕｎｄｅｒ　ｔｈｅ　ｄｅｐｏｓｉｔｓ　ｏｆ　ｔｅｒｔｉａｒｙ　ｃｅｌｌ　ｗａｌｌｓ．　Ｔｈｅ　ｄｅｐｏｓｉｔｉｏｎ　ｏｆ　ｓｕｂｅｒｉｎ　ｉｓｏｌａｔｅｓ　ｔｈｅ　ｅｎｄｏｄｅｒｍａｌ　ｃｅｌｌ　ｐｒｏｔｏｐｌａｓｔ
ｆｒｏｍ　ｔｈｅ　ａｐｏｐｌａｓｔ，　ｔｈｕｓ　ｒｅｔａｒｄｉｎｇ　Ｃａ２＋　ｉｎｆｌｕｘ　ｔｏ　ｅｎｄｏｄｅｒｍａｌ　ｃｅｌｌｓ　ａｎｄ　ｔｈｅｒｅｂｙ　ｃａｌｃｉｕｍ　ｔｒａｎｓｐｏｒｔ　ｔｏ　ｔｈｅ　ｘｙｌｅｍ．　Ｉｎ　ｔｈｅ　ｃｅｌｌ　ｏｆ　ｒｏｏｔ　ｔｉｐ，
Ｃａｓｐａｒｉａｎ　ｂａｎｄ　ｈａｓ　ｎｏｔ　ｄｅｖｅｌｏｐｅｄ　ｙｅｔ，　ａｎｄ　ｃａｌｃｉｕｍ　ｉｓ　ａｌｌｏｗｅｄ　ｔｏ　ｔｒａｖｅｒｓｅ　ｔｈｅ　ｅｎｄｏｄｅｒｍｉｓ　ｖｉａ　ｔｈｅ　ａｐｏｐｌａｓｔｉｃ　ｐａｔｈｗａｙ．　Ｗｈｅｎ　ｃａｌｃｉｕｍ
ｇｅｔｓ　ｔｏ　ｔｈｅ　ｓｔｅｌａｒ　ａｐｏｐｌａｓｔ，　ｉｔ　ｃａｎ　ｅｎｔｅｒ　ｘｙｌｅｍ　ｖｅｓｓｅｌ　ｔｈｒｏｕｇｈ　Ｃａ２＋　ｃｈａｎｎｅｌｓ　ｉｎ　ｔｈｅ　ａｃｃｅｓｓｉｂｌｅ　ｐｌａｓｍａ　ｍｅｍｂｒａｎｅ　ｏｆ　ｔｈｅ　ｘｙｌｅｍ　ｐａｒｅｎｃｈｙｍａ
ｃｅｌｌｓ，　ｗｈｉｃｈ　ｈａｖｅ　ｉｏｎ　ｃｈａｎｎｅｌｓ　（Ｔｅｓｔｅｒ　ａｎｄ　Ｌｅｉｇｈ　２００１）．　Ｏｎｌｙ　ａ　ｌｉｔｔｌｅ　ｃａｌｃｉｕｍ　ｉｓ　ｔｒａｎｓｆｅｒｒｅｄ　ｔｏ　ｔｈｅ　ｘｙｌｅｍ　ｖｉａ　ｔｈｅ　ｓｙｍｐｌａｓｔｉｃ　ｐａｔｈｗａｙ．

Casparian band has not been formed and the endoder-
mal Casparian band may lose its function as an
apoplastic barrier when roots undergo secondary
(thickening) growth (Peterson and Lefcourt 1990). The
functioning of the apoplastic barriers is not limited to
any single mode. Rather, it changes with the changes
in chemical composition of the barriers, and therefore
can be regulated through changing their composition
(Hose1 et al. 2001).
2.2 Transport from epidermal cell to xylem in root
When the Ca2+ moving forward to xylem reaches
endodermis through the apoplastic pathway, the bar-
rier formed of Casparian strip forces it to enter the
root symplast through Ca2+ channels in the accessible
plasma membrane of the endodermal cells (Fig.1). It
is then actively effused from the symplast by the plasma
membrane Ca2+-ATPase or Ca2+/H+ antiporter of cells
within the stele. To reach the xylem, Ca2+ transported
230 　植物生理与分子生物学学报
into endodermal cell must subsequently be secreted
into the stelar apoplast. A corollary of this is that two
protein-mediated steps are necessary for the delivery
of solutes to the xylem, one for uptake into the sym-
plast (whether at the epidermis, cortex or endodermis)
and the other for release from stelar cells. There is a
steeper Ca 2+ gradient from cell wall to cytoplasm,
which drives Ca 2+ flux to symplast through Ca 2 +
channels, but the process that Ca2+ effuses from sym-
plast and then release to xylem from pericycle stelar
parenchyma needs some pump to drive. Electrophysi-
ological studies confirm the presence of transport pro-
teins in the plasma membrane of trichoblasts, which is
needed for the uptake of Ca2+ (Tester and Leigh 2001).
There is a considerable electrochemical gradient
to drive the Ca2+ influx through Ca2+ channels in any
root cell (White 1998; Kiegle et al. 2000). In proto-
plasts isolated from the xylem parenchyma of steles of
barley roots, three types of cation-selective channels
have been identified, which makes it possible for cat-
ion to be released from the xylem parenchyma into the
xylem apoplast through ion channels (Wegner and
Raschke 1994). Several types of Ca2+-permeable chan-
nels in the plasma membrane of root cells have been
recorded. By using electrophysiological techniques, the
presence of Ca2+-permeable channels has been dem-
onstrated (White 1998; Véry and Sentenac 2002),
which is likely to be responsible for the loading of sol-
utes into the xylem. Patch clamping of protoplasts iso-
lated from the xylem parenchyma of barley and the
stele of Arabidopsis showed clearly the presence of
both cation and anion channels with characteristics
consistent with a role in the loading of solutes into
xylem (Tester and Leigh 2001).
Ca2+-ATPase is one kind of Ca2+ pump that drives
2+
Ca flux from symplast to apoplast. The presence of
Ca2+-ATPase in the plasma membrane of root cells has
been proved both biochemically (Evans and Williams
1998; Giesler et al. 2000) and electrophysiologically
(Felle et al. 1992).
The distribution of Ca2+-ATPase has relation to
the direction Ca2+ of transport. In apple root cells, Ca2+-
ATPase is distributed primarily in the plasmalemma,
the inner margin of vessels and the space between plas-
malemma and cell wall, and its distribution is not uni-
form in cell and plasmalemma. The activity of Ca2+ -
　３１ 卷
ATPases decreases gradually in radial direction from
vessel to epidermis cells (Yang et al. 2004). This indi-
cates that it is feasible for this transport mechanism to
pump Ca2+ to the stelar apoplast.
In Arabidopsis, the SOS1 gene encoding the pu-
tative Na+/H+ antiporter preferentially is expressed at
the xylem symplast boundary of roots (indicated by
promoter-GUS fusion) (Shi et al. 2002). At moderate
Na+ concentrations, SOS1 is proposed to function in
the active efflux of Na+ to the xylem, because sos1
mutants accumulated less Na+ in shoot than did the wild
type. Since Ca2+ /H+ antiporter can transport Ca2+ to
the vacuole, it should be able to drive Ca2+ flux from
symplast to apoplast, but no evidence of such flux has
been found at the xylem symplast boundary of roots.
The control of xylem loading of ion by ABA is rel-
evant to these considerations. ABA can inhibit the chan-
nel-mediated efflux of K+ and Cl– into xylem (Roberts
1998), but stimulate H+ extrusion into the xylem, a pro-
cess that tends to stimulate any Na+/H+ antiporter ac-
tivity in the plasma membrane of xylem parenchyma
cells (Tester and Davenport 2003). There have been
few works on the function of Ca2+/H+ antiporter in the
loading of Ca2+ in xylem.
2.3 Transport in trunk xylem
After being released to xylem, Ca2+ is transported
to the aerial part through mass flow of water and ion
exchange. Water flow is what drives Ca2+ flow, so their
rates are highly correlated in the transport to shoot via
xylem (Quintana et al. 1999). In orange xylem, Ca2 +
can move up and down and be transferred between
phloem and xylem. The Ca2+ being transported in plant
xylem vessel is in chelate form, such as calcium cit-
rate or calcium malate. The presence of organic acid
can impact the activity of cation, which may be impor-
tant for Ca2+ transport in orange xylem (Xiao and Hu
1997). In root xylem sap of mature beech (Fagus
sylvatica L.) trees, high Ca2+ and malic acid concen-
trations were observed when the pH value was low,
and negative correlation between them was established.
These high correlations suggest that the malic acid
content determines the low pH value of the xylem sap
and malic acid tends to form complexes with hydrated
or exchangeably adsorbed cations, thereby accelerates
the mobilization and translocation of Ca 2+ (Schell
1997).
３期 杨洪强等：　植物钙素吸收和运转（英文）
Movement of Ca2+-containing xylem sap is driven
by two processes: growth-induced water influx and
transpiration. For the higher rate, transpiration is the
main factor of driving Ca2+ movement. Low rates of
transpiration can lead to low rates of Ca2+ accretion
into aerial organs. In snap bean, variability of pod Ca2+
concentration among genotypes is due to differences
in pod transpiration rate (Grusak and Pomper 1999;
Quintana et al. 1999). Wounding may alter the flow of
xylem sap. After a brief scorching of tomato leaves
with a blowlamp, the leaves release water into the trunk,
which carries a pulse of up to 50% of the leaf’s total
calcium into the trunk and makes xylem Ca2+ levels
increase (Malone et al. 2002).
3 Transport in leaves and fruits
Evidence shows that there is a barrier on the path
of water and solutes moving free from shoot xylem to
leaf. The first is that there is that tracer of apoplastic
flow accumulates in the bundle sheath apoplast and
the solute potential is different between the xylem and
mesophyll apoplast. The second is that the relation
between solute movement through leaf and transpira-
tional flow is not simple. Ca2+ moves from leaf vascu-
lar tissue to the epidermis via vein extensions. The
selectivity of cell to ion is different, so there is more
Ca 2+ a ccumulates in the e pidermis than in the
mesophyll. Discrimination against Ca2+ uptake into the
symplast at the plasma membrane of the bundle sheath
cells prevents Ca2+ accumulation in the mesophyll
(Karley et al. 2000).
The movement of potassium is easier than cal-
cium in general, but calcium uptake is faster than po-
tassium in the first 60 d of Actinidia deliciosa fruit
development that lasts about 150 d. On the 60th day
after fruit set, only about 40% of the time of fruit de-
velopment has passed, the calcium content has reached
about 70% of that at harvest, whereas potassium
reached only 50%. This high value of fruit calcium
uptake was correlated with high fruit transpiration
observed during the first 60 days after fruit set. After
this period, the fruit transpiration declined and fruit
calcium concentration and Ca/K ratio decreased for
fruit growth rapidly, while calcium concentration in-
creased constantly in leaves. This behaviour confirms
231
that calcium movement between leaf and fruit through
the phloem is limited (Xiloyannis et al. 2001).
Rate of Ca2+ accumulation in “Golden Delicious”
apple is higher during the first half of the fruit-grow-
ing season than in the second half. Spraying of 0.15%
Ca2+ solution every two weeks during the first half of
the fruit growing season did not make the calcium con-
tent any higher. During the second half, treated fruits
accumulated more Ca2+ and final Ca2+ contents were
higher. This suggests that there is no absorption of Ca2+
after it is being sprayed on fruits during the first grow-
ing period, for the Ca2+ is mainly provided by root ab-
sorption (Casero et al. 2002). But the results of ex-
periment in high-density apple orchard show that the
total calcium content in fruit increases and calcium
concentration was significantly higher in the peel than
in the flesh at harvest after frequent canopy sprays of
Ca2+ on fruit from the time of fruit set to the end of
Augus t (Zavalloni et al. 2001). The re are both
apoplastic and symplastic pathways of Ca2+ transport
in fruit tissue, the apoplastic pathway is the main one
in apple (Zhou and Lin 2000). The transport through
apoplastic pathway relies mainly on transpiration and
that through symplastic pathway relies on Ca2+-ATPase
that drives Ca2+ to move into the fruit (Zhou and He
1999).
4 Perspectives
Ca2+ absorption by cells is a passive process be-
cause many inhibitors of respiration which inhibit the
entering of phosphorus and potassium into cells do not
inhibit that of calcium (Shear and Faust 1970), so it
has been believed that the amount of Ca2+ absorb and
transport through the whole plant depends only on the
rate of flow of transpiration stream. But recent results
of research show that the rate of Ca2+ uptake by root is
higher at proper temperature and is inhibited by the
metabolic inhibitor 2,4-dinitrophenol (Yang et al.
2003), which indicates that plant Ca2+ uptake is not
solely a passive process. The absorption of Ca2+ by
the whole plant is more complicated than that by a
cell. At whole plant level, the uptake and transport in-
volve many cells and traverse through symplastic and
apoplastic pathways, which consist of three main steps:
traversing the root cortex to xylem, transporting in
232 　植物生理与分子生物学学报
xylem and distributing to leaves or fruits. The results
of the study on cell level cannot be used directly to
explain the mechanism of Ca2+ uptake by whole plant
and the methods used in the study at cell level cannot
be used directly the study at whole plant level. Some
special methods to study the transport of Ca2+ at whole
plant level are needed.
Movement in xylem proceeds only through the
apoplast. In going into and out of xylem, Ca2+ may
traverse through either an apoplastic or symplastic
pathway, or both. But the relative contribution of them
is not known yet. The mechanism of Ca2+ release to
xylem vessel from xylem parenchyma cell is not clear
yet.
Casparian band is the key position of ion trans-
port root. When Ca2+ reaches there when traversing
the root cortex apoplastically, it must enter the sym-
plast of the endodermal cell through Ca2+ channels
before being actively fluxed to the stele through Ca2+-
ATPases. There are reports showing that the rate of
Ca2+ uptake in apple root increases with an increase in
Ca2+-ATPase activity. IBA can increase Ca2+-ATPase
activity and the rate of Ca2+ uptake in apple root (Yang
et al. 2000, 2004). These observations indicate Ca2+-
ATPase is important for Ca2+ uptake in root. New re-
port shows that IBA can activate Ca2+-ATPase activity
of Malus hupehensis roots through protein phospho-
rylation (Li et al. 2004). Protein kinase catalyzes pro-
tein phosphorylation. Both protein kinase and Ca2+ play
important roles in plant signaling (Reddy 2001; Yang
and Liang 2001). But the role of protein phosphoryla-
tion in the uptake and transport of nutrient element is
not clear yet. The activity Ca2+-ATPase can be regu-
lated by phosphorylation, so the role of phosphoryla-
tion in Ca2+ uptake should be an important part for
research in the future.
The uptake and accumulation of Ca2+ in pepper
fruits are different significantly from the plant species
(Tang et al. 2002), which indicates that it is possible
to select the efficient genotype of Ca2+ using as root-
stock or variety. Application of functional genomics
may provide insights to Ca2+ uptake and accumulation
in further study.
Ｒｅｆｅｒｅｎｃｅｓ
Ｃａｓｅｒｏ　Ｔ，　Ｂｅｎａｖｉｄｅｓ　Ａ，　Ｒｅｃａｓｅｎｓ　Ｉ，　Ｒｕｆａｔ　Ｊ　（２００２）．　Ｐｒｅｈａｒｖｅｓｔ
　３１ 卷
Ｃａ　ｓｐｒａｙｓ　ａｎｄ　ｆｒｕｉｔ　Ｃａ　ａｂｓｏｒｐｔｉｏｎ　ｉｎ　ｇｏｌｄｅｎ　 ‘ａｐｐｌｅｓ’
Ａｃｔａ　Ｈｏｒｔ，　５ ９ ４：　４６７－４７３
Ｄａｖｅｎｐｏｒｔ　Ｒ，　Ｔｅｓｔｅｒ　Ｍ　（２０００）．　Ａ　ｗｅａｋｌｙ　ｖｏｌｔａｇｅ－ｄｅｐｅｎｄｅｎｔ，
ｎｏｎ－ｓｅｌｅｃｔｉｖｅ　ｃａｔｉｏｎ　ｃｈａｎｎｅｌ　ｍｅｄｉａｔｅｓ　ｔｏｘｉｃ　ｓｏｄｉｕｍ　ｉｎｆｌｕｘ
ｉｎ　ｗｈｅａｔ．　Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ，　１２２：　８２３－８３４
ｄｅ　Ｂｏｅｒ　ＡＨ　（１９９９）．　Ｐｏｔａｓｓｉｕｍ　ｔｒａｎｓｌｏｃａｔｉｏｎ　ｉｎｔｏ　ｔｈｅ　ｒｏｏｔ　ｘｙｌｅｍ．
Ｐｌａｎｔ　Ｂｉｏｌ，　１：　３６－４５
Ｅｖａｎｓ　ＤＥ，　Ｗｉｌｌｉａｍｓ　ＬＥ　（１９９８）．　Ｐ－ｔｙｐｅ　ｃａｌｃｉｕｍ　ＡＴＰａｓｅｓ　ｉｎ
ｈｉｇｈｅｒ　ｐｌａｎｔｓ—bｉｏｃｈｅｍｉｃａｌ，　ｍｏｌｅｃｕｌａｒ　ａｎｄ　ｆｕｎｃｔｉｏｎａｌ
ｐｒｏｐｅｒｔｉｅｓ．　Ｂｉｏｃｈｉｍ　Ｂｉｏｐｈｙｓ　Ａｃｔａ，　１ ３ ７ ６：　１－２５
Ｆｅｌｌｅ　ＨＨ，　Ｔｒｅｔｙｎ　Ａ，　Ｗａｇｎｅｒ　Ｇ　（１９９２）．　Ｔｈｅ　ｒｏｌｅ　ｏｆ　ｔｈｅ　ｐｌａｓｍａ
ｍｅｍｂｒａｎｅ　Ｃａ２＋－ＡＴＰａｓｅ　ｉｎ　Ｃａ２＋　ｈｏｍｅｏｓｔａｓｉｓ　ｉｎ　Ｓｉｎａｐｉｓ
ａｌｂａ　ｒｏｏｔ　ｈａｉｒｓ．　Ｐｌａｎｔａ，　１８８：　３０６－３１３
Ｇｉｅｓｌｅｒ　Ｍ，　Ａｘｅｌｓｅｎ　ＫＢ，　Ｈａｒｐｅｒ　ＪＦ，　Ｐａｌｍｇｒｅｎ　ＭＧ　（２０００）．
Ｍｏｌｅｃｕｌａｒ　ａｓｐｅｃｔｓ　ｏｆ　ｈｉｇｈｅｒ　ｐｌａｎｔ　Ｐ－ｔｙｐｅ　Ｃａ２＋－ＡＴＰａｓｅｓ．
Ｂｉｏｃｈｉｍ　Ｂｉｏｐｈｙｓ　Ａｃｔａ，　１４６５ ：　５２－７８
Ｇｉｌｒｏｙ　Ｓ，　Ｊｏｎｅ　ＤＬ　（２０００）．　Ｔｈｒｏｕｇｈ　ｆｏｒｍ　ｔｏ　ｆｕｎｃｔｉｏｎ：　ｒｏｏｔ　ｈａｉｒ
ｄｅｖｅｌｏｐｍｅｎｔ　ａｎｄ　ｎｕｔｒｉｅｎｔ　ｕｐｔａｋｅ．　Ｔｒｅｎｄｓ　Ｐｌａｎｔ　Ｓｃｉ，　５（２）：
５６－６０
Ｇｒｕｓａｋ　ＭＡ，　Ｐｏｍｐｅｒ　ＫＷ　（１９９９）．　Ｉｎｆｌｕｅｎｃｅ　ｏｆ　ｐｏｄ　ｓｔｏｍａｔａｌ
ｄｅｎｓｉｔｙ　ａｎｄ　ｐｏｄ　ｔｒａｎｓｐｉｒａｔｉｏｎ　ｏｎ　ｔｈｅ　ｃａｌｃｉｕｍ　ｃｏｎｃｅｎｔｒａｔｉｏｎ
ｏｆ　ｓｎａｐ　ｂｅａｎ　ｐｏｄｓ．　Ｊ　Ａｍ　Ｓｏｃ　Ｈｏｒｔ　Ｓｃｉ，　１ ２ ４：　１９４－１９８
Ｈｏｓｅ１　Ｅ，　Ｃｌａｒｋｓｏｎ　ＤＴ，　Ｓｔｅｕｄｌｅ　Ｅ，　Ｓｃｈｒｅｉｂｅｒ　Ｌ，　Ｈａｒｔｕｎｇ　Ｗ
（２００１）．　Ｔｈｅ　ｅｘｏｄｅｒｍｉｓ：　ａ　ｖａｒｉａｂｌｅ　ａｐｏｐｌａｓｔｉｃ　ｂａｒｒｉｅｒ．　Ｊ
Ｅｘｐｔ　Ｂｏｔ，　５２：　２２４５－２２６４
Ｋａｒｌｅｙ　ＡＪ，　Ｌｅｉｇｈ　ＲＡ，　Ｓａｎｄｅｒｓ　Ｄ　（２０００）．　Ｗｈｅｒｅ　ｄｏ　ａｌｌ　ｔｈｅ　ｉｏｎｓ
ｇｏ？　Ｔｈｅ　ｃｅｌｌｕｌａｒ　ｂａｓｉｓ　ｏｆ　ｄｉｆｆｅｒｅｎｔｉａｌ　ｉｏｎ　ａｃｃｕｍｕｌａｔｉｏｎ　ｉｎ
ｌｅａｆ　ｃｅｌｌｓ．　Ｔｒｅｎｄｓ　Ｐｌａｎｔ　Ｓｃｉ，　５（１１）：　４６５－４７０
Ｋｉｅｇｌｅ　Ｅ，　Ｇｉｌｌｉｈａｍ　Ｍ，　Ｈａｓｅｌｏｆｆ　Ｊ，　Ｔｅｓｔｅｒ　Ｍ　（２０００）．
Ｈｙｐｅｒｐｏｌａｒｉｓａｔｉｏｎ－ａｃｔｉｖａｔｅｄ　ｃａｌｃｉｕｍ　ｃｕｒｒｅｎｔｓ　ｆｏｕｎｄ　ｏｎｌｙ
ｉｎ　ｃｅｌｌｓ　ｆｒｏｍ　ｔｈｅ　ｅｌｏｎｇａｔｉｏｎ　ｚｏｎｅ　ｏｆ　Ａｒａｂｉｄｏｐｓｉｓ　ｔｈａｌｉａｎａ
ｒｏｏｔｓ．　Ｐｌａｎｔ　Ｊ，　２１：　２２５－２３１
Ｌｅｎｇ　ＱＲＷ，　Ｍｅｒｃｉｅｒ　ＷＹ，　Ｂｅｒｋｏｗｉｔｚ　ＧＡ　（１９９９）．　Ｃｌｏｎｉｎｇ　ａｎｄ
ｆｉｒｓｔ　ｆｕｎｃｔｉｏｎａｌ　ｃｈａｒａｃｔｅｒｉｚａｔｉｏｎ　ｏｆ　ａ　ｐｌａｎｔ　ｃｙｃｌｉｃ
ｎｕｃｌｅｏｔｉｄｅ－ｇａｔｅｄ　ｃａｔｉｏｎ　ｃｈａｎｎｅｌ．　Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ，　１ ２１：　７５３－
７６１
Ｌｉ　Ｊ（李佳），　Ｙａｎｇ　ＨＱ（杨洪强），　Ｙａｎ　Ｂ（闫滨），　Ｓｈｕ　ＨＲ（束怀瑞）
（２００４）．　Ｉｎｄｏｌｅ　ｂｕｔｙｒｉｃ　ａｃｉｄ　ａｃｔｉｖａｔｅｓ　Ｃａ２＋－ＡＴＰａｓｅ　ｉｎ　Ｍａｌｕｓ
ｈｕｐｅｈｅｎｓｉｓ　Ｒｅｈｄ．　ｒｏｏｔｓ　ｂｙ　ｐｒｏｔｅｉｎ　ｐｈｏｓｐｈｏｒｙｌａｔｉｏｎ．　Ｊ
Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ　Ｍｏｌ　Ｂｉｏｌ　（植物生理与分子生物学学报），
３０ （４）：　４４９－４５４　（ｉｎ　Ｃｈｉｎｅｓｅ）
Ｍａａｔｈｕｉｓ　ＦＪＭ，　Ｍａｙ　ＳＴ，　Ｇｒａｈａｍ　ＮＳ，　Ｂｏｗｅｎ　ＨＣ，　Ｊｅｌｉｔｔｏ　ＴＣ，
Ｔｒｉｍｍｅｒ　Ｐ，　Ｂｅｎｎｅｔｔ　ＭＪ，　Ｓａｎｄｅｒｓ　Ｄ，　Ｗｈｉｔｅ　ＰＪ　（１９９８）．　Ｃｅｌｌ
ｍａｒｋｉｎｇ　ｉｎ　Ａｒａｂｉｄｏｐｓｉｓ　ｔｈａｌｉａｎａ　ａｎｄ　ｉｔｓ　ａｐｐｌｉｃａｔｉｏｎ　ｔｏ
ｐａｔｃｈ－ｃｌａｍｐ　ｓｔｕｄｉｅｓ．　Ｐｌａｎｔ　Ｊ，　１５ ：　８４３－８５１
Ｍａｌｏｎｅ　Ｍ，　Ｗｈｉｔｅ　ＰＪ，　Ａｎｇｅｌａ　ＭＭ　（２００２）．　Ｍｏｂｉｌｉｚａｔｉｏｎ　ｏｆ
ｃａｌｃｉｕｍ　ｉｎ　ｇｌａｓｓｈｏｕｓｅ　ｔｏｍａｔｏ　ｐｌａｎｔｓ　ｂｙ　ｌｏｃａｌｉｚｅｄ
ｓｃｏｒｃｈｉｎｇ．　Ｊ　Ｅｘｐ　Ｂｏｔ，　５３ ：　８３－８８
Ｐａｕｌ　ＣＢ，　Ｒｕｓｓｅｌ　ＬＪ　（２０００）．　Ｖａｃｕｏｌｅｓ　ａｎｄ　ｐｒｅｖａｃｕｏｌａｒ
ｃｏｍｐａｒｔｍｅｎｔｓ．　Ｃｕｒｒ　Ｏｐｉｎ　Ｐｌａｎｔ　Ｂｉｏｌ，　３ （６）：　４６９－４７５
Ｐｅｔｅｒｓｏｎ　ＣＡ，　Ｃｈｏｌｅｗａ　Ｅ　（１９９８）．　Ｓｔｒｕｃｔｕｒａｌ　ｍｏｄｉｆｉｃａｔｉｏｎｓ　ｏｆ
３期 杨洪强等：　植物钙素吸收和运转（英文）
ｔｈｅ　ａｐｏｐｌａｓｔ　ａｎｄ　ｔｈｅｉｒ　ｐｏｔｅｎｔｉａｌ　ｉｍｐａｃｔ　ｏｎ　ｉｏｎ　ｕｐｔａｋｅ．　Ｊ
Ｐｌａｎｔ　Ｎｕｔｒ　Ｓｏｉｌ　Ｓｃｉ，　１６１：　５２１－５３１
Ｐｅｔｅｒｓｏｎ　ＣＡ，　Ｌｅｆｃｏｕｒｔ　ＢＥＭ　（１９９０）．　Ｄｅｖｅｌｏｐｍｅｎｔ　ｏｆ
ｅｎｄｏｄｅｒｍａｌ　Ｃａｓｐａｒｉａｎ　ｂａｎｄｓ　ａｎｄ　ｘｙｌｅｍ　ｉｎ　ｌａｔｅｒａｌ　ｒｏｏｔｓ
ｏｆ　ｂｒｏａｄ　ｂｅａｎ．　Ｃａｎ　Ｊ　Ｂｏｔ，　６８ ：　２７２９－２７３５
Ｑｕｉｎｔａｎａ　ＪＭ，　Ｈａｒｒｉｓｏｎ　ＨＣ，　Ｐａｌｔａ　ＪＰ，　Ｎｉｅｎｈｕｉｓ　Ｊ，　Ｋｍｉｅｃｉｋ　Ｋ
（１９９９）．　Ｘｙｌｅｍ　ｆｌｏｗ　ｒａｔｅ　ｄｉｆｆｅｒｅｎｃｅｓ　ａｒｅ　ａｓｓｏｃｉａｔｅｄ　ｗｉｔｈ
ｇｅｎｅｔｉｃ　ｖａｒｉａｔｉｏｎ　ｉｎ　ｓｎａｐ　ｂｅａｎ　ｐｏｄ　ｃａｌｃｉｕｍ　ｃｏｎｃｅｎｔｒａｔｉｏｎ．
Ｊ　Ａｍ　Ｓｏｃ　Ｈｏｒｔ　Ｓｃｉ，　１２４ （５）：　４８８－４９１
Ｒｅｄｄｙ　ＡＳＮ　（２００１）．　Ｃａｌｃｉｕｍ：　Ｓｉｌｖｅｒ　ｂｕｌｌｅｔ　ｉｎ　ｓｉｇｎａｌｉｎｇ． 　Ｐｌａｎｔ
Ｓｃｉ，　１ ６ ０：　３８１－４０４
Ｒｏｂｅｒｔｓ　ＳＫ　（１９９８）．　Ｒｅｇｕｌａｔｉｏｎ　ｏｆ　Ｋ ＋　ｃｈａｎｎｅｌｓ　ｉｎ　ｍａｉｚｅ　ｒｏｏｔｓ
ｂｙ　ｗａｔｅｒ　ｓｔｒｅｓｓ　ａｎｄ　ＡＢＡ．　Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ，　１１６：　１４５－１５３
Ｓｃｈｅｌｌ　Ｊ　（１９９７）．　Ｉｎｔｅｒｄｅｐｅｎｄｅｎｃｅ　ｏｆ　ｐＨ，　ｍａｌａｔｅ　ｃｏｎｃｅｎｔｒａｔｉｏｎ，
ａｎｄ　ｃａｌｃｉｕｍ　ａｎｄ　ｍａｇｎｅｓｉｕｍ　ｃｏｎｃｅｎｔｒａｔｉｏｎｓ　ｉｎ　ｔｈｅ　ｘｙｌｅｍ
ｓａｐ　ｏｆ　ｂｅｅｃｈ　ｒｏｏｔｓ．　Ｔｒｅｅ　Ｐｈｙｓｉｏｌ，　１７：　４７９－４８３
Ｓｈｅａｒ　ＲＬ，　Ｆａｕｓｔ　Ｍ　（１９７０）．　Ｃａｌｃｉｕｍ　ｔｒａｎｓｐｏｒｔ　ｉｎ　ａｐｐｌｅ　ｔｒｅｅｓ．
Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ，　１５：　６７０－６７４
Ｓｈｅａｒ　ＣＢ　（１９７５）．　Ｃａｌｃｉｕｍ－ｒｅｌａｔｅｄ　ｄｉｓｏｒｄｅｒｓ　ｏｆ　ｆｒｕｉｔｓ　ａｎｄ
ｖｅｇｅｔａｂｌｅｓ．　ＨｏｒｔＳｃｉｅｎｃｅ　１０：　３６１－３６５
Ｓｈｉ　Ｈ，　Ｑｕｉｎｔｅｒｏ　ＦＪ，　Ｐａｒｄｏ　ＪＭ，　Ｚｈｕ　ＪＫ　（２００２）．　Ｔｈｅ　ｐｕｔａｔｉｖｅ
ｐｌａｓｍａ　ｍｅｍｂｒａｎｅ　Ｎａ＋／Ｈ＋　ａｎｔｉｐｏｒｔｅｒ　ＳＯＳ１　ｃｏｎｔｒｏｌｓ　ｌｏｎｇ－
ｄｉｓｔａｎｃｅ　Ｎａ＋　ｔｒａｎｓｐｏｒｔ　ｉｎ　ｐｌａｎｔｓ．　Ｐｌａｎｔ　Ｃｅｌｌ，　１４：　４６５－４７７
Ｔａｎｇ　ＷＨ（唐文浩），　Ｃｈｅｎ　ＸＨ（陈雪华），　Ｋｕａｎｇ　ＣＬ（邝春兰），　Ｘｉａ
ＦＪ（夏福君），　Ｙａｎｇ　Ｐ（杨萍）　（２００２）．　Ｇｅｎｏｔｙｐｉｃ　ｄｉｆｆｅｒｅｎｃｅｓ
ａｎｄ　ｐｈｙｓｉｏｌｏｇｉｃａｌ　ｃｈａｒａｃｔｅｒｉｓｔｉｃｓ　ｏｆ　ｃａｌｃｉｕｍ　ｕｐｔａｋｅ　ａｎｄ
ａｃｃｕｍｕｌａｔｉｏｎ　ｏｆ　ｐｅｐｐｅｒ　（Ｃａｐｓｉｃｕｍ　ｆｒｕｔｅｓｃｅｎｓ）．　Ｐｌａｎｔ　Ｎｕｔｒ
Ｆｅｒｔｉｌｉｚｅｒ　Ｓｃｉ　（植物营养与肥料学报），　８（３）：　３４９－３５４　（ｉｎ
Ｃｈｉｎｅｓｅ）
Ｔｅｓｔｅｒ　Ｍ，　Ｄａｖｅｎｐｏｒｔ　Ｒ　（２００３）．　Ｎａ＋　ｔｏｌｅｒａｎｃｅ　ａｎｄ　Ｎａ＋　ｔｒａｎｓｐｏｒｔ
ｉｎ　ｈｉｇｈｅｒ　ｐｌａｎｔｓ．　Ａｎｎ　Ｂｏｔ，　９１（５）：　５０３－５２７
Ｔｅｓｔｅｒ　Ｍ，　Ｌｅｉｇｈ　ＲＡ　（２００１）．　Ｐａｒｔｉｔｉｏｎｉｎｇ　ｏｆ　ｎｕｔｒｉｅｎｔ　ｔｒａｎｓｐｏｒｔ
ｐｒｏｃｅｓｓｅｓ　ｉｎ　ｒｏｏｔｓ．　Ｊ　Ｅｘｐ　Ｂｏｔ，　５２：　４４５－４５７
Ｕｅｏｋａ－Ｎａｋａｎｉｓｈｉ　Ｈ，　Ｎａｋａｎｉｓｈｉ　Ｙ，　Ｔａｎａｋａ　Ｙ，　Ｍａｅｓｈｉｍａ　Ｍ
（１９９９）．　Ｐｒｏｐｅｒｔｉｅｓ　ａｎｄ　ｍｏｌｅｃｕｌａｒ　ｃｌｏｎｉｎｇ　ｏｆ　Ｃａ２＋／Ｈ＋
ａｎｔｉｐｏｒｔｅｒ　ｉｎ　ｔｈｅ　ｖａｃｕｏｌａｒ　ｍｅｍｂｒａｎｅ　ｏｆ　ｍｕｎｇ　ｂｅａｎ．　Ｅｕｒｏ
Ｊ　Ｂｉｏｃｈｅｍ，　２ ６ ２：　４１７－４２５
Ｖérｙ　ＡＡ，　Ｄａｖｉｅｓ　ＪＭ　（２０００）．　Ｈｙｐｅｒｐｏｌａｒｉｚａｔｉｏｎ－ａｃｔｉｖａｔｅｄ
ｃａｌｃｉｕｍ　ｃｈａｎｎｅｌｓ　ａｔ　ｔｈｅ　ｔｉｐ　ｏｆ　Ａｒａｂｉｄｏｐｓｉｓ　ｒｏｏｔ　ｈａｉｒｓ．　Ｐｒｏｃ
Ｎａｔｌ　Ａｃａｄ　Ｓｃｉ　ＵＳＡ，　９７ 　（１７）：　９８０１－９８０６
Ｖérｙ　ＡＡ，　Ｓｅｎｔｅｎａｃ　Ｈ　（２００２）．　Ｃａｔｉｏｎ　ｃｈａｎｎｅｌｓ　ｉｎ　ｔｈｅ　Ａｒａｂｉｄｏｐｓｉｓ
ｐｌａｓｍａ　ｍｅｍｂｒａｎｅ．　Ｔｒｅｎｄｓ　Ｐｌａｎｔ　Ｓｃｉ，　７（４）：　１６８－１７５
Ｗｅｇｎｅｒ　ＬＨ，　Ｒａｓｃｈｋｅ　Ｋ　（１９９４）．　Ｉｏｎ　ｃｈａｎｎｅｌｓ　ｉｎ　ｔｈｅ　ｘｙｌｅｍ
ｐａｒｅｎｃｈｙｍａ　ｏｆ　ｂａｒｌｅｙ　ｒｏｏｔｓ：　ａ　ｐｒｏｃｅｄｕｒｅ　ｔｏ　ｉｓｏｌａｔｅ
ｐｒｏｔｏｐｌａｓｔｓ　ｆｒｏｍ　ｔｈｉｓ　ｔｉｓｓｕｅ　ａｎｄ　ａ　ｐａｔｃｈ－ｃｌａｍｐ　ｅｘｐｌｏｒａｔｉｏｎ
233
ｏｆ　ｓａｌｔ　ｐａｓｓａｇｅｗａｙｓ　ｉｎｔｏ　ｘｙｌｅｍ　ｖｅｓｓｅｌｓ．　Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ，
１０５ ：　７９９－８１３
Ｗｈｉｔｅ　ＰＪ　（２００１）．　Ｔｈｅ　ｐａｔｈｗａｙｓ　ｏｆ　ｃａｌｃｉｕｍ　ｍｏｖｅｍｅｎｔ　ｔｏ　ｔｈｅ
ｘｙｌｅｍ．　Ｊ　Ｅｘｐ　Ｂｏｔ，　５２ ：　８９１－８９９
Ｗｈｉｔｅ　ＰＪ　（２０００）．　Ｃａｌｃｉｕｍ　ｃｈａｎｎｅｌｓ　ｉｎ　ｈｉｇｈｅｒ　ｐｌａｎｔｓ．　Ｂｉｏｃｈｉｍ
Ｂｉｏｐｈｙｓ　Ａｃｔａ，　１４６５ ：　１７１－１８９
Ｗｈｉｔｅ　ＰＪ　（１９９８）．　Ｃａｌｃｉｕｍ　ｃｈａｎｎｅｌｓ　ｉｎ　ｔｈｅ　ｐｌａｓｍａ　ｍｅｍｂｒａｎｅ
ｏｆ　ｒｏｏｔ　ｃｅｌｌｓ．　Ａｎｎ　Ｂｏｔ，　８１ ：　１７３－１８３
Ｗｈｉｔｅ　ＰＪ，　Ｂｒｏａｄｌｅｙ　ＭＲ　（２００３）．　Ｃａｌｃｉｕｍ　ｉｎ　ｐｌａｎｔｓ．　Ａｎｎ　Ｂｏｔ，　９２
（４）：　４８７－５１１
Ｘｉａｏ　ＬＴ（萧浪涛），　Ｈｕ　ＤＪ（胡笃敬）　（１９９７）　．　Ｔｈｅ　ｕｐｔａｋｅ　ａｎｄ　ｆｌｕｘ
ａｎａｌｙｓｉｓ　ｏｆ　Ｃａ２＋　ｉｎ　ｃｉｔｒｕｓ　ｒｏｏｔ．　Ａｃｔａ　Ｐｈｙｔｏｐｈｙｓｉｏｌ　Ｓｉｎ　（植
物生理学报），　２３（３）：　２３３－２３８　（ｉｎ　Ｃｈｉｎｅｓｅ）
Ｘｉｌｏｙａｎｎｉｓ　Ｇ，　Ｃｅｌａｎｏ　Ｇ，　Ｍｏｎｔａｎａｒｏ　Ｂ，　Ｄｉｃｈｉｏ　Ｌ，　Ｓｅｂａｓｔｉａｎｉ
ＡＭ　（２００１）．　Ｗａｔｅｒ　ｒｅｌａｔｉｏｎｓ，　ｃａｌｃｉｕｍ　ａｎｄ　ｐｏｔａｓｓｉｕｍ
ｃｏｎｃｅｎｔｒａｔｉｏｎ　ｉｎ　ｆｒｕｉｔｓ　ａｎｄ　ｌｅａｖｅｓ　ｄｕｒｉｎｇ　ａｎｎｕａｌ　ｇｒｏｗｔｈ
ｉｎ　ｍａｔｕｒｅ　ｋｉｗｉｆｒｕｉｔ　ｐｌａｎｔｓ．　Ａｃｔａ　Ｈｏｒｔ，　５６４：　１２９－１３４
Ｙａｎｇ　ＨＱ（杨洪强），　Ｊｉｅ　ＹＬ（接玉玲），　Ｚｈａｎｇ　ＬＺ（张连忠）　（２０００）．
Ｅｆｆｅｃｔ　ｏｆ　ＩＢＡ　ｏｎ　ｔｈｅ　Ｃａ２＋　ｕｐｔａｋｅ　ａｎｄ　ｒｏｏｔ　ａｃｔｉｖｉｔｙ　ｉｎ　ｙｏｕｎｇ
ａｐｐｌｅ　ｔｒｅｅｓ．　Ａｄｖ　Ｈｏｒｔ　（园艺学进展），　４：　５７－５９　（ｉｎ　Ｃｈｉｎｅｓｅ）
Ｙａｎｇ　ＨＱ　（ 杨洪强），　Ｚｈａｎｇ　ＬＺ　（ 张连忠），　Ｑｉ　ＪＬ（戚金亮），　Ｊｉｅ
ＹＬ（接玉玲）　（２００３）．　Ｔｈｅ　ｋｉｎｅｔｉｃｓ　ｏｆ　ｃａｌｃｉｕｍ　ｕｐｔａｋｅ　ｉｎ
ａｐｐｌｅ　ｒｏｏｔｓｔｏｃｋ　ｒｏｏｔｓ．　Ａｃｔａ　Ｈｏｒｔ　Ｓｉｎ　（园艺学报），　３０（３）：
２５３－２５６　（ｉｎ　Ｃｈｉｎｅｓｅ）
Ｙａｎｇ　ＨＱ，　Ｊｉｅ　ＹＬ，　Ｚｈａｎｇ　ＬＺ，　Ｃｕｉ　ＭＧ．　（２００４）．　Ｔｈｅ　ｅｆｆｅｃｔ　ｏｆ　ＩＢＡ
ｏｎ　ｔｈｅ　Ｃａ２＋　ａｂｓｏｒｐｔｉｏｎ　ａｎｄ　Ｃａ２＋－ＡＴＰａｓｅ　ａｃｔｉｖｉｔｙ　ａｎｄ　ｔｈｅｉｒ
ｕｌｔｒａｃｙｔｏｃｈｅｍｉｃａｌ　ｌｏｃａｌｉｚａｔｉｏｎ　ｉｎ　ａｐｐｌｅ　ｒｏｏｔｓ．　Ａｃｔａ　Ｈｏｒｔ，
６３６ ：　２１１－２１９
Ｙａｎｇ　ＨＱ（杨洪强），　Ｌｉａｎｇ　ＸＥ（梁小娥）　（２００１）． Ｐｒｏｔｅｉｎ　ｋｉｎａｓｅ
ａｎｄ　ｅｎｖｉｒｏｎｍｅｎｔａｌ　ｓｔｒｅｓｓ　ｓｉｇｎａｌｉｎｇ　ｃａｓｃａｄｅｓ　ｉｎ　ｐｌａｎｔｓ．
Ｐｌａｎｔ　Ｐｈｙｓｉｏｌ　Ｃｏｍｍｕｎ（植物生理学通讯），　３ ７ 　（３）：　１８５－
１９４　（ｉｎ　Ｃｈｉｎｅｓｅ）
Ｚａｖａｌｌｏｎｉ　Ｃ，　Ｍａｒａｎｇｏｎｉ　Ｂ，　Ｔａｇｌｉａｖｉｎｉ　Ｍ，　Ｓｃｕｄｅｌｌａｒｉ　Ｄ　（２００１）．
Ｄｙｎａｍｉｃｓ　ｏｆ　ｕｐｔａｋｅ　ｏｆ　ｃａｌｃｉｕｍ，　ｐｏｔａｓｓｉｕｍ　ａｎｄ　ｍａｇｎｅｓｉｕｍ
ｉｎｔｏ　ａｐｐｌｅ　ｆｒｕｉｔ　ｉｎ　ａ　ｈｉｇｈ　ｄｅｎｓｉｔｙ　ｐｌａｎｔｉｎｇ．　Ａｃｔａ　Ｈｏｒｔ，　５ ６４：
１１３－１２１
Ｚｈｏｕ　Ｗ　（周卫），　Ｈｅ　Ｐ（何萍）（１９９９）．　Ｔｈｅ　ｃｈａｒａｃｔｅｒｉｓｔｉｃ　ｏｆ　Ｃａ２＋－
ＡＴＰａｓｅ　ａｃｔｉｖａｔｅｄ　ｔｒａｎｓｐｏｒｔ　ｏｆ　Ｃａ２＋　ｉｎ　ｔｈｅ　ｐｌａｓｍａｌｅｍｍａ　ｏｆ
ａｐｐｌｅ　ｆｒｅｓｈ．　Ａｃｔａ　Ｐｈｙｔｏｐｈｙｓｉｏｌ　Ｓｉｎ　（植物生理学报），　２５
（２）：　１５１－１５８　（ｉｎ　Ｃｈｉｎｅｓｅ）
Ｚｈｏｕ　Ｗ（周卫），　Ｌｉｎ　Ｐ（林葆）　（２０００）．　Ｓｔｕｄｙ　ｏｎ　ｐａｔｈｗａｙｓ　ｏｆ　Ｃａ２＋
ｍｏｖｅｍｅｎｔ　ｉｎ　ｙｏｕｎｇ　ｆｒｕｉｔ　ｔｉｓｓｕｅ　ｏｆ　ａｐｐｌｅ　ａｎｄ　ｉｔｓ　ｒｅｇｕｌａｔｉｏｎ
ｂｙ　ｈｏｒｍｏｎｅｓ．　Ｐｌａｎｔ　Ｎｕｔｒ　Ｆｅｒｔｉｌｉｚｅｒ　Ｓｃｉ　（植物营养与肥料
学报），　６（２）：　２１４－２１９　（ｉｎ　Ｃｈｉｎｅｓｅ）
 234 植物生理与分子生物学学报，　Ｊｏｕｒｎａｌ　 ｏ ｆ　 Ｐ ｌａｎｔ　 Ｐ ｈｙｓｉｏｌｏｇｙ　 ａ ｎｄ　 Ｍ ｏｌｅｃｕｌａｒ　 Ｂ ｉｏｌｏｇｙ　　 ２ ００５，　 ３ １　 （ ３）：　 ２ ２７ －２３４

植物钙素吸收和运转
杨洪强 ＊，接玉玲
（山东农业大学园艺学院，泰安　２７１０１８）

摘要：近年来，钙素在植物体内的吸收和运输研 能需要 Ｃａ ２＋ －ＡＴＰ 酶驱动；还有一些 Ｃａ ２＋ 由内皮层

究主要集中在细胞和分子水平，但整株水平上的 细胞运出，沿内皮层内侧的质外体途径进入木质
研究也同样重要。整株水平上的钙吸收和运输包 部导管，并通过导管运向枝干。钙离子以螯合态
括根细胞的钙吸收、钙离子横向穿过根系并进入 的形式在枝干导管运输；水流速率是影响钙离子
木质部、在木质部运输、从木质部移出并进入叶 沿导管运输的关键因子。钙离子在果实和叶片中
片或果实及在叶片或果实中运转分配等环节，既 的运输和分配不仅通过质外体途径也通过共质体途
２＋
经过质外体也穿越共质体。钙离子通道、Ｃ ａ － 径。
ＡＴＰ 酶和 Ｃａ ２＋ ／Ｈ ＋ 反向转运器等参与根细胞的钙吸
收。在钙离子横向穿根进入木质部的过程中，需 关键词：钙；根系；吸收运转；Ｃ ａ ２ ＋ － Ａ Ｔ Ｐ 酶；木质部
要穿越内皮层和木质部薄壁细胞组织。根系内皮 中图分类号：Ｑ９４５．１ ；Ｓ１４３．７

层凯氏带阻挡了 Ｃａ ２ ＋ 沿质外体途径由内皮层外侧
向内侧的移动，部分 Ｃａ ２ ＋ 由此通过离子通道流进
内皮层细胞而转入共质体并到达木质部薄壁细胞组 国家自然科学基金项目（Ｎｏｓ．　３０１７０６５５， ３０２７０９２３）资助。
＊Ｅ－ｍａｉｌ：　ｌａｂｆｔ＠ｓｄａｕ．ｅｄｕ．ｃｎ；　ｙｈｑｓｄ＠ｙａｈｏｏ．ｃｏｍ．ｃｎ
织，而由木质部薄壁细胞组织进入中柱质外体可

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