Long-Distance Transport

in the Xylem and Phloem

Mochammad Roviq, 2011
 • The long-distance transport of water and
solutes - mineral elements and low-
Concept molecularweight organic compounds - takes
place in the vascular system of xylem and
phloem.
• Long-distance transport from the roots to the
shoots occurs predominantly in the nonliving
xylem vessels.
• Coniferous trees lack the continuous system
of xylem vessels, and depend on tracheides
which are non-living conducting cells ranging
in length from 2 to 6 mm.
• In annual plant species too long-distance
transport in the xylem vessels may be
interrupted by tracheides, for example at the
root-shoot junction or in the nodes of the
stem.
• These structures pose an internal resistance
to xylem volume flow but simultaneously
permit an intensive xylem phloem solute
transfer
Root pressure
• Xylem transport is driven by the
gradient in hydrostatic pressure
(root pressure) and by the gradient
in the water potential
• The gradient in water potential
between roots and shoots is quite
steep particularly during the day
when the stomata are open.
• Values become less negative in the
following sequence: atmosphere »
leaf cells > xylem sap > root cells >
external solution
Direction of long-distance
transport
• Solute flow in the xylem from the
roots to the shoots is therefore
unidirectional
• Under certain conditions in the shoots
a counterflow of water in the xylem
may also occur, for example, from
low-transpiring fruits back to the
leaves
• Long-distance transport in the phloem
takes place in the living sieve tube
cells and is bidirectional.
• The direction of transport is
determined primarily by the
nutritional requirements of the
various plant organs or tissues and
occurs, therefore, from source to sink
Transfer cells
• During long-distance
transport, mineral
elements and organic
solutes are transferred
between the xylem and
phloem by extensive
exchange processes,
referred to as loading and
unloading.
• The transfer is mediated by
specific cells called transfer
cells
Transfer cells are specialised
parenchymal cells.
• They are different from
the other cells of the
plant structure in that,
they are concerned with
transport of solutes.
• They are present in high
numbers wherever
active absorption and
secretion are evident.
• They are mostly found in
the close conjunction
with vascular cambial
cells i.e., xylem and
phloem
 Composition of the Xylem Sap
• Depend on factors such as plant species, mineral
element supply to the roots, assimilation of
mineral nutrients in the roots and nutrient
recycling.
• Composition and particularly concentration of
solutes are also strongly influenced by the degree
of dilution by water and are therefore dependent
on the transpiration rate and the time of day.
• Composition and concentration of xylem sap also
change typically during the ontogenesis of plants
Xylem Volume Flow (Pressurized Exudation at 100 kPa) and Mineral
Element Concentrations in the Xylem Sap of Soil-Grown Nodulated
Soybean During Reproductive Stage
 Exchange Adsorption and Resorption
• The interactions between cations are similar to those in
the AFS of the root cortex .
• The degree of retardation (penghambatan) of cation
translocation depends on the valency of the cation (Ca2+>
K+), its own activity, the activity of competing cations, the
charge density of the negative groups (dicots > monocots),
the pH of the xylem sap which may vary between 5 and 7
and the diameter of the xylem vessels.
• The translocation rate in the xylem of heavy metal cations
is much enhanced when the ions are complexed, for
example, in the case of copper, zinc or cadmium
 Resorption
• Solutes can also be resorbed from the xylem
(apoplasm) into the living cells (transport in the
cytoplasm and vacuole) along the pathway of the
xylem sap from the roots to the leaves
• Resorption from the xylem can be the result either of
transient or permanent storage in the xylem
parenchyma and other stem tissue, or of xylem-
phloem transfer, mediated by specialized cells (xylem
parenchyma transfer cells)
• The resorption of certain mineral elements from the
xylem sap is very pronounced and can have important
consequences for the mineral nutrition of these plants
 Natrophobic or natrophilic plant species

• Na+ is retained mainly in the roots and lower

stem (Phaseolus vulgaris), whereas in natrophilic
species (e.g. sugar beet) translocation into the
leaves readily occurs
 Resorspsi unsur mikro
• Resorpsi dari cairan xilem di akar dan batang juga dapat
menjadi faktor penentu dalam distribusi unsur mikro pada
tanaman.
• Sehingga ketika pasokan
molibdenum dalam
medium hara tinggi, pada
tomat terjadi keracunan
jauh lebih awal daripada
kacang atau bunga
matahari
 Release or Secretion
• Komposisi cairan xilem di sepanjang
jalur transportasi juga dapat berubah
oleh pelepasan atau sekresi zat terlarut
dari sel-sel di sekitarnya.
• Misalnya, nonlegum yang diberi nitrat,
konsentrasi nitrat di cairan xilem
menurun sejalan dengan semakin
panjang jalur, sedangkan konsentrasi
nitrogen organik, khususnya glutamin
meningkat
• Dalam legum yang bernodulasi (di mana
terjadi fiksasi N2), di sisi lain, rasio
amida dengan asam amino bergeser
mendukung asam amino
 Xylem Unloading in Leaves
• Meskipun terjadi resorpsi sepanjang jalur di batang, sebagian
besar zat terlarut dan air diangkut pembuluh xilem ke daun.
• Air diangkut dalam pembuluh utama ke bagian yang
mengalami penguapan cepat seperti tepi daun
• Konsentrasi zat terlarut
dapat naik dibagian
tertentu, misalnya, tepi
daun, tergantung pada
konsentrasi dan komposisi
larutan dalam cairan xilem
yang masuk daun, laju
kehilangan air melalui
transpirasi, dan panjang
jalur yang melalui daun
Precipitation…..
• Pencegahan akumulasi zat terlarut yang
berlebihan dalam apoplasm daun dapat
dicapai dengan pembentukan garam low
solubility di apoplasm, terutama untuk
menghilangkan kalsium larut dalam
gymnosperms
• This mechanism of precipitation
seems to be a safe way of coping with
a continuous xylem import of calcium Calcium oxalate crystals in the
which is scarcely exported in the apoplasm of needles. (Left) Micrograph
from the phloem of a needle from
phloem and where the ionic Juniperus chinensis; (right) micrograph
of a stomatal pore of a needle from
concentrations in the symplasm have Picea abies (L.) Karst.
to be kept very low.
 Model for scavenging solutes from the xylem sap
(xylem unloading') in leaf cells.
• The cells of the bundle
sheath are sites of intensive
net proton excretion which
acidifies the apoplasm the
proton gradient across the
plasma membrane acting as
the driving force for
cotransport of amino acids
and ureides.
• The activity of the proton
pump in legumes is high
before pod filling and
disappears during pod As- amino acids.
formation
 Effect of Transpiration
Rate on Uptake and
Translocation

• Tingkat fluks air di akar

A. 'Passive' transport of mineral elements through
the apoplasm into the stele. B. More rapid removal
of mineral elements released into the xylem vessels
. C. Increase in the mass flow of the external
solution to the rhizoplane and eventually into the
apparent free space of the cortex, favoring active
uptake into the symplasm. E, Endodermis; X, xylem;
arrow, water flux (A to C see text).
(transportasi jarak pendek) dan
di dalam pembuluh xilem
(transportasi jarak jauh)
ditentukan oleh tekanan akar
dan laju transpirasi.
Peningkatan tingkat transpirasi
mungkin, atau tidak mungkin,
meningkatkan serapan dan
translokasi unsur mineral dalam
xilem tersebut. Peningkatan
dapat dicapai dengan berbagai
cara
Predominantly factors of uptake and
translocation rate:
1. Plant age. In seedlings and young plants with
a low leaf surface area, enhancement effects
of transpiration are usually absent; water
uptake and solute transport in the xylem to
the shoots are determined mainly by the root
pressure. As the age and size of the plants
increase, the relative importance of the
transpiration rate, particularly for the
translocation of mineral elements increases.
 2. Time of day
• In leaves up to 90% of the total transpiration is
stomatal. During the light period, transpiration rates
and thus the potential enhancement of uptake and
translocation of mineral elements are much higher
than during the dark period.
translocation rate

• 3. External Effect of Transpiration Rate of Sugar Beet Plants

concentration. on Uptake and Translocation of Potassium and
Sodium from Nutrient Solutions'
increase in the
concentration of
mineral elements
in the nutrient
medium may
enhance the effect
of transpiration
rate on the uptake
and translocation
of mineral
elements.
translocation rate

• 4. Type of mineral element. Under otherwise

comparable conditions (e.g. plant age and
external concentration), the effect of
transpiration rates on the uptake and transport
follows a typically defined ranking order of
mineral elements.
• transpirasi meningkatkan penyerapan dan
translokasi molekul tak bermuatan ke tingkat
yang lebih besar daripada ion.
• Ada hubungan erat antara tingkat transpirasi dan
tingkat penyerapan herbisida tertentu
 Measured and Calculated Silicon Uptake in
Relation to Transpiration (Water Consumption)
of Oat Plants
 Effect of Transpiration Rate on
Distribution within the Shoot
• The long-distance transport of a mineral element
exclusively in the xylem should be expected to
give a distinct distribution pattern in the shoot
organs that depends on both transpiration rates
(e.g., ml g-1 dry weight each day) and duration of
transpiration (e.g. age of the organ).
• For example, manganese (where at the same
plant (maple tree) and similar leaf age the *sun
leaves' (high transpiration rates) have much
higher manganese contents in their dry matter
than 'shade leaves' (low transpiration rates).
 Increasing supply…

• The distribution of boron is also

related to the loss of water
from the shoot organ, as shown
by the boron distribution in
shoots of rape in response to
an increasing boron supply
• The typical gradient in the
Effect of increasing boron
transpiration rates in the shoot application to the soil on the
organs (leaves > pods » seeds) distribution of boron in the
corresponds to the gradient in shoots of rape

boron content.
 II. Phloem Transport
• Principles of Transport and Phloem Anatomy
• Composition of the Phloem Sap
• Mobility in the Phloem
• Transfer between the Xylem and Phloem
 Principles of Transport and Phloem
Anatomy

• Long-distance transport in
the phloem takes place in
living cells, the sieve
tubes
• The principles of the
transport mechanism in
the phloem were
proposed as early as 1930
by Munch in pressure
flow hypothesis
(Druckstromtheorie) Cross-sectional area of a vascular bundle from
based on the principle of the stem of maize. Inset: sieve tube with sieve
the osmometer. plate pores and T-protein'. (From Eschrich,
1976.)
 Release or unloading at the sink
• Munch suggested that solutes
such as sucrose are
concentrated in the phloem of
leaves (i.e., phloem loading) and
the water is sucked into the
phloem, creating a positive
internal pressure.
• This pressure induces a mass
flow in the phloem to the sites
of lower positive pressure
caused by removal of solutes
from the phloem.
• Flow rate and direction of flow
are therefore closely related to
the release or unloading at the
sink.
Munch pressure flow hypothesis
• This type of pressure-
driven mass flow in the
phloem differs from that
in the xylem in three
important ways: (a)
Organic compounds are
the dominant solutes in
the phloem sap; (b)
transport takes place in
living cells; and (c) the
unloading of solutes at
the sink plays an
important role.
 Source-sink regulated transport
• An example for a primarily
source-sink regulated transport
of a mineral nutrient is shown
in for phosphorus.
• After application to one of the
two mature primary leaves,
the labelled phosphorus is
transported to the shoot apex
and the roots whereas
transport to the other primary
leaf is negligible.
• In contrast, sodium is not Retranslocation of labeled phosphorus
transported to the shoot apex (P) and sodium (Na) after application to
but exclusively moves the tip of a primary leaf of bean.
downwards (basipetally) to the Autoradiogram, 24 h after application.
 Composition of the Phloem Sap
• Phloem sap has a high pH (7-8) and contains high
concentrations of solutes, on average 15-25% dry matter.
• The main component is usually sucrose, which may
comprise up to 90% of the solids. The proportion of sucrose
to other solutes depends on the site of phloem sap
collection, it is very high, for example near the ear of
cereals
• In addition to sucrose, among the other organic solutes
amino acids are usually present in high concentrations the
amides glutamine and asparagine may represent up to 90%
of this fraction, whereas the concentrations of nitrate and
ammonium are usually very low
Comparison of the Levels of Organic and Inorganic Solutes in the
Phloem and Xylem Exudates of Nicotiana glauca
 Mobility in the Phloem

• The classification is, of

course, only a first
approximation as certain
factors are ignored, for
example, genotypical
differences or the nutritional
status of plants.
• For the macronutrients,
except calcium (phloem
mobility is generally high,
and for the micronutrients it
is at least intermediate with
the exception of manganese.
 Transfer between the Xylem and
Phloem

• In the vascular
bundles, phloem and
xylem are separated
by only a few cells
(Lihat gambar
sebelumnya).
• In the regulation of
long-distance
transport, exchange
of solutes between
the two conducting
systems is very
important
Water and solute flow
 Transfer from xylem to phloem
• Dari perbedaan konsentrasi yang ditampilkan di
dalamnya adalah jelas bahwa transfer dari floem
ke xilem dapat terjadi downhill, melalui membran
plasma dari tabung saringan, jika sebuah gradien
konsentrasi yang cukup ada.
• Sebaliknya, bagi sebagian besar zat terlarut
organik dan anorganik ditransfer dari xilem ke
floem biasanya merupakan uphill transport
melawan gradien konsentrasi antara apoplasm
(xilem) dan symplasm sel xilem sekitar parenkim
dan sel-sel floem
 Higher demand

• The xylem-to-phloem transfer is of

• This transfer of organic and
Long-distance transport in xylem (X)
and phloem (P) in a stem with a
connected leaf, and xylem-to-phloem
transfer mediated by a transfer cell (T).
particular importance for the
mineral nutrition of plants,
because xylem transport is
directed mainly to the sites
(organs) of highest transpiration,
which are usually not the sites of
highest demand for mineral
nutrients.
inorganic solutes can take place all
along the pathway from roots to
shoot, and the stem plays an
important role in this respect most
likely via transfer cells
 Mineral Nutrients with High Phloem
Mobility

• For mineral nutrients with

high phloem mobility such
as potassium, phosphorus
or nitrogen as amino-N the
relative importance of
phloem and xylem
transport into an organ
mainly depends on the
stage of development of
the organ as shown in for
amino-N during the life of
an individual leaf.
 Mineral Nutrients of Low Phloem
Mobility; Example Calcium

• Because of its low concentrations in the phloem sap (Section 3.3.2)

the import of calcium into growth sinks such as shoot apices, young
leaves or fruits takes place nearly exclusively in the xylem, whereas
the import in the phloem is negligible as shown for castor bean in
Table 3.11. This is in marked contrast to potassium of which most
(terminal bud) and at least half (youngest leaves) of the total net
import takes place in the phloem. For magnesium phloem import
contributes to 25 and 40% of the total import, respectively.
 Environmental Factors
• To increase the
calcium content
of growing leaves
or fruits,
increasing the
transpiration
rates of the fruits
is more effective
than increasing
the calcium
supply in the Effect of Environmental Factors and Growth Rate
on Calcium and Potassium Content of Red Pepper
substrate Fruits
 Root pressure
• High root pressure, as indicated by the intensity
of guttation, is closely correlated with an
increased concentration of calcium in expanding
leaves and either the absence of, or only mild
symptoms of, calcium deficiency (tip necrosis).
• Magnesium, which is highly phloem mobile, is
only slightly affected by root pressure.
• Root pressure also strongly depends on root
respiration and oxygen supply to the roots.
Guttation
Retranslocation and Cycling of
Nutrients
• With exception of calcium and presumably also
manganese, import of nutrients in the xylem and
export (retranslocation) in the phloem is a normal
feature throughout the life of an individual leaf.
• Several pieces of evidence indicate a rapid xylem-
to-phloem transfer in the leaf blades and
involvement of only a small fraction of the total
leaf content ('cycling' fraction;) in this process.
• Between 82 and 100% of the
exported mineral elements
are retranslocated in the
phloem back to the roots, and
a high proportion of the
potassium and magnesium
cycle, i.e., they are again
loaded into the xylem and
transported to the shoots.
• For calcium and sodium no
precise data can be given but
cycling is of minor
importance. For cycling of
potassium, corresponding
data for other plant species
are 20% in tomato and 30% in
wheat and rye.
 Remobilization of Mineral Nutrients
• Import and export of mineral nutrients occur
simultaneously during the life of plant organs
such as leaves (Table 3.16).
 Physiological and biochemical
processes
• Remobilization is based on a range of different
physiological and biochemical processes:
– utilization of mineral nutrients stored in vacuoles
(potassium, phosphorus, magnesium, amino-N, etc.),
– breakdown of storage proteins (e.g., in vacuoles of the
paraveinal mesophyll cells of legumes;), or, finally,
– breakdown of cell structures (e.g., chloroplasts) and
– enzyme proteins thereby transforming structurally
bound mineral nutrients (e.g., magnesium in
chlorophyll, micronutrients in enzymes) into a mobile
form.
 Seed Germination
• During the germination of seeds (or storage
organs such as tubers) mineral nutrients are
remobilized within the seed tissue and
translocated in the phloem or xylem, or both,
to the developing roots or shoots.
• As a rule, seedlings will grow for at least
several days without an external supply of
mineral nutrients
 Vegetative Stage
• During vegetative growth, nutrient supply to the roots is
often either permanently insufficient (as in the case of low
soil nutrient content) or temporarily interrupted (when, for
example, there is a lack or excess of soil moisture).
• Remobilization of mineral nutrients from mature leaves to
areas of new growth is thus of key importance for the
completion of the life cycle of plants under these
experimental conditions.
• This behaviour (strategy) is typical for fast-growing crop
species whereas for many wild species cessation of growth
occurs under adverse environmental conditions and, thus,
redistribution of mineral nutrients plays a lesser important
role
 Reproductive Stage
• Remobilization of mineral nutrients is particularly
important during reproductive growth when
seeds, fruits, and storage organs are formed.
• At this growth stage root activity and nutrient
uptake generally decrease, mainly as a result of
decreasing carbohydrate supply to the roots ('sink
competition'.
• Therefore, the mineral nutrient contents of
vegetative parts quite often decline sharply
during the reproductive stage
 Period Before Leaf Drop (Perennials)
• As a rule, and similar to annual species, the
extent of remobilization is high for nitrogen,
potassium, phosphorus, and zinc, whereas the
leaf contents of calcium, boron, iron and
manganese increase until leaf drop
• Remobilization of mineral nutrients (except
calcium and manganese) from the leaves to
woody parts is a typical feature of perennial
species before leaf drop in temperate climates,
and is closely related to the discoloration of
leaves in the autumn.
Terimakasih