Annals of Botany 83 : 19–27, 1999

Article No. anbo.1998.0781, available on line at http:\\www.idealibrary.com

Water Retention Capacity in Root Segments Differing in the Degree of

Exodermis Development
E. T A L E I S N IK*, G. P E Y R A NO, A. C O! R D O BA and C. A R I A S
Instituto de FitopatologıT a y FisiologıT a Vegetal, INTA, Cna 60 Cuadras Km 5 1\2, CP 5119, CoT rdoba, Argentina

Received : 15 June 1998 Returned for revision : 5 August 1998 Accepted : 10 September 1998

Water loss from roots back into drying soil is a problem of practical importance in plants growing under conditions
of very low substrate water potential, such as dry or saline areas. Root exodermis is relatively impermeable and has
been suggested to play a protective role against water loss. The relative water retention ability was compared in root
segments from exodermal (maize, onion, sunflower, Rhodes grass and sorghum) and non-exodermal species (Pisum
satiŠum, Vicia faba and wheat). Apical and basal segments from exodermal roots, with different degrees of exodermis
development, were also compared, as were segments from sorghum roots in which the exodermis thickness had been
modified by subjecting the plants to a 30 d water stress treatment. Water retention was significantly higher in segments
from exodermal roots. In each root, water loss was higher in apical than in basal segments, regardless of the presence
of exodermis. In sorghum, prolonged drought treatment increased exodermis thickening in nodal roots, however, no
differences in rates of water loss were observed in segments obtained from control and droughted plants. Soil sheaths
formed around roots of Rhodes grass growing in very dry soil with the epidermis adhering tightly to the sheath. In
plants growing in the field, soil sheaths may be more effective than the exodermis in preventing root water loss.
# 1999 Annals of Botany Company
Key words : Root, exodermis, rhizosheaths, water loss.

suberized layers were associated with decreasing radial

INTRODUCTION
Water loss from roots backing into drying soil is a problem
for plants growing under conditions of restricted water
supply or very low substrate water potential, such as saline
areas. However, available information regarding this issue
is controversial (Passioura, 1988). In some species, signifi-
cant water loss from the root to the substrate has been
reported. Water was passively lost along a water potential
gradient from roots to drying soil by bermudagrass
(Cynodon dactyloniCynodon transŠaalensis L. Pers.) (Baker
and van Bavel, 1986), wheat (Blum and Johnson, 1992) and
pearl millet (Vetterlein and Marschner, 1993).
The mechanisms of water transport in and across roots
have been extensively reviewed (see Boyer, 1985 ; Passioura,
1988 ; Oertli, 1996). Radial water transport in roots occurs
along apoplasmic and cell-to-cell routes (Steudle, 1995) ;
these pathways have different relative importance in different
species (Steudle, Murrmann and Peterson, 1993) and present
various resistances. The main apoplasmic resistances are the
exodermis and endodermis which form the outer and inner
boundaries of the root cortex, respectively. The exodermis is
a hypodermis with suberized walls and Casparian bands
(Henberg, 1968) ; it derives, like the cortex, from the ground
meristem, can be uni- or multiseriate and is internal to the
epidermis. The exodermis becomes fully developed at
varying distances from the root tip (Clarkson et al., 1987),
depending on the species. In general, available data indicate
that suberin lamellae in the exodermis of a root reduce the
permeability to water (Peterson and Enstone, 1996). These
* For correspondence. Fax 54 51 974330
0305-7364\99\010019j09 $30.00\0
hydraulic conductance along root axes (Melchior and
Steudle, 1993) and are a barrier to the diffusion of
apoplasmic dyes (Enstone and Peterson, 1992).
Environmental stresses such as drought and salinity can
decrease root hydraulic conductance (Munns and Passioura,
1984 ; Peyrano et al., 1997) and increase exodermis
suberization. In sorghum, the thickness of this layer
increased with decreasing water availability (Cruz, Jordan
and Drew, 1992) and this was suggested to have a bearing
on the observed reduction in root radial hydraulic con-
ductance. In cotton, exposure to high salinity (200 m
NaCl) induced the formation of an exodermis with
Casparian bands and suberin lamellae close to the root base
and in the transition zone to the hypocotyl (Reinhardt and
Rost, 1995). The exodermis, which did not develop in
control roots, was suggested to play a role in protecting the
root from water loss and\or leakage of solutes important
for osmotic adjustment. Under water deficit conditions,
death of the root cortex may occur (Jupp and Newman,
1987), however, it is retained in exodermal roots (Peterson,
1989), and it was suggested that cells with suberized walls
arrest dieback in roots by apparently reducing water loss
from the root to the soil.
Studies with roots and isolated sleeves of root epidermal\
exodermal cells of maize, suggest that the suberized walls
remain relatively permeable when bathed by culture sol-
ution, but become impermeable if exposed to moist air
rather than being kept completely immersed in culture
solution (Clarkson et al., 1987). Under such conditions, the
exodermis would impose a significant resistance to water
loss. From the suggested role of the exodermis in restricting
# 1999 Annals of Botany Company
20 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots
water loss from roots it may be inferred that plants with
exodermal roots have a relative advantage in dry or saline
environments. This study, therefore, examines the relative
water retention ability of exodermal and non-exodermal
roots. It compares rates of water loss in segments from
exodermal and non-exodermal roots, in apical and basal
segments from exodermal roots (which have a different
degree of exodermis development) and in sorgum roots in
which the degree of exodermis thickness was modified by
subjecting the plants to a prolonged water stress period.
MATERIALS AND METHODS
Roots from species which differ in the degree of exodermal
development were used. Species with a developed exodermis
were maize (Zea mays L.), onion (Allium cepa L.), sunflower
(Helianthus annus L.), Rhodes grass (Chloris gayana Kunth)
and sorghum (Sorghum bicolor (L.) Moench). Species
without exodermis development were pea (Pisum satiŠum
L.), Vicia faba L. and wheat (Triticum aestiŠum L.)
(Perumalla, Peterson and Enstone, 1990).
Plants were grown in a glasshouse, in pots with vermiculite
irrigated with half-strength Hoagland solution. In the case
of sorghum, 2 month old plants growing in vermiculite were
subjected to a 1 month water stress treatment in which the
substrate in control plants was kept moist by frequent
irrigation while droughted plants were irrigated once a week
and only if the upper 3 cm of vermiculite were visibly dry.
Root sections for the study were obtained from nodal roots.
In addition, roots from C. gayana plants growing in dry soil
in the province of Santiago del Estero, Argentina, were
observed for soil sheath formation.
Root segments, 20 mm in length, were obtained from two
zones : apical (comprising the root tip, which was cut off)
and basal (approx. 100 mm from the apex). The rate of
water loss in such segments was measured by two methods :
(1) by assessing the time required to reach equilibrium
within the sample chamber of a psychrometer ; or (2) by
periodically determining weight loss in samples allowed to
dry on the lab bench under room conditions. Temperature
and relative humidity were registered in transpiration
measurements. In both cases, segments were sealed at both
ends with silicon grease to prevent water loss from the cut
ends.
Free-hand sections were obtained from segments similar
to the ones used for water loss measurements using a
modification of Fro$ hlich’s technique (Fro$ hlich, 1984).
Sections were placed in holder chambers and stained
essentially as described by Brundrett, Enstone and Peterson
(1988), replacing toluidine blue for aniline blue (O’Brien,
Feder and McCully, 1964). Sections were observed in a
Zeiss Axiophot microscope with UV illumination using
excitation filter 395–440, chromatic beam splitter FT 460
and barrier filter LP 470. Unstained sections were observed
for autofluorescence under the same setting. Photographs
were taken with Kodak Ektachrome slide film ASA 400.
Exodermal wall thickness was measured in the photographs
using the Optimas 6.1 program for image analysis.
Since roots of different diameters differ in their surface
area : volume ratio, it was reasoned that thinner roots, with
a larger surface area : volume ratio, would lose water at a
faster rate. To test this hypothesis, water loss was measured
in agar cylinders of different diameters. These were obtained
by allowing 12 % agar solutions to gel within capillary tubes
having internal diameters in the range of the studied root
segments. Agar cylinders were ejected with the aid of an air-
filled syringe, weighed immediately and then sequentially to
determine the rate of water loss when exposed to air under
room conditions.
RESULTS
Water loss from exodermal and non-exodermal root
segments
It was expected that root segments differing in the degree of
exodermal development would take different lengths of time
to reach equilibrium in a psychrometer chamber (Kikuta
and Richter, 1992). However, the required equilibrium time
was similar for apical and basal segments from the same
root (Fig. 1), irrespective of whether species were exodermal
or not (Fig. 2 A and B). Fifth order polynomials gave a good
fit for the range of points in those curves (R#
0n95). Pair-
wise comparisons of coefficients for the different powers of
x from apical and basal segments were performed for each
species, and probability values for the t tests indicated
curves for apical and basal segments were not significantly
different for each species.
Psychrometer results suggested there was no difference in
rates of water loss between root segments with different
degrees of exodermis development. These results were
verified by periodically weighing root segments exposed to
air. The rate of water loss was initially faster in segments
from non-exodermal species (Fig. 3), and, after 20 min,
when the relative water content was approx. 75 %, water
retention had become significantly lower in segments from
non-exodermal roots.
In maize, a species with exodermal roots, water loss from
apical segments (where the exodermis is not yet suberized),
was faster than in basal segments (with suberized exodermis),
after 10 min of air exposure (Fig. 4). However, after 20 min
water loss was significantly faster in apical than basal
segments (Table 1) in roots of peas that do not develop a
suberized exodermis (Fig. 2 A).
Water loss as a function of root diameter
Apical segments are tapered and thinner than basal
segments, and differences in rates of water loss could
therefore be the result of different surface area : volume
ratios (Cape and Percy, 1996). In 12 % agar cylinders of
different diameters, weight retention was proportional to
diameter (Fig. 5), indicating the rate of water loss was
inversely proportional to the thickness of the cylinder.
However, in root segments, root diameter and weight
retention were not significantly correlated (correlation
coefficient l 0n24, R# l 0n06, Fig. 6). Weight retention after
30 min of air exposure was higher in exodermal roots, as
seen from the distribution of points above the line indicating
50 % retention (Fig. 6).
 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots 21
50

40

30
µVolt

20

10

0
40

30
µVolt

20

10

0 10 20 30 40 50 60 70
Time (min)
F. 1. Voltage readings in a psychrometer cell for root segments of exodermal ($, #) and non-exodermal roots (
, ) obtained between
5–13 mm (apical ; #, ) and 100–108 mm (basal ; $,
) from the root apex. Results are from a single root in each case.

In sorghum nodal roots, a prolonged drought treatment

Effect of salinity and drought on exodermis deŠelopment
and root water loss
In Chloris gayana, root hydraulic conductance decreases
after 30 d treatment with 300 m NaCl (Peyrano, unpubl.
res.). This prompted us to examine whether decreased
hydraulic conductance was associated with changes in
exodermis wall thickness, and whether these would affect
root water loss.
Water potential in control and salinized roots was
0n19p0n07 and 1n72p0n04 MPa, but increased development
of the exodermis was not detected in salinized plants (Fig.
2 C and D). After 30 min of air exposure, water retention in
basal segments from salinized roots was less than in control
roots (Fig. 7).
Soil sheaths formed around roots of C. gayana growing in
very dry soil (Fig. 2 G). In transverse sections of such roots,
the epidermis and exodermis were seen to adhere tightly to
the soil sheath, and were separated from the stelar tissues by
a lacunar cortex (Fig. 2 H).
increased exodermis thickness (Fig. 2 F and G). Tangential
cell walls in the exodermis of droughted roots were 4n78 µm
thick, 1n62-times thicker than controls (t lk7n09, 16 d.f., P
0n001). However, no differences in rates of water loss were
observed in basal segments obtained from control and
droughted plants (Fig. 8).
DISCUSSION
Exodermal root segments were expected to take longer than
non-exodermal segments to reach equilibrium within psy-
chrometer cells (Kikuta and Richter, 1992). However,
equilibration times were similar for both exodermal and
non-exodermal segments. This is understandable if equi-
librium is reached with water loss from the outermost
epidermal layer, and is unaffected by features that could
differentially affect water loss. Nevertheless, after 20 min of
exposure to air, water retention in exodermal segments was
significantly higher than in non-exodermal segments ; this
22 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots

a e
r

F. 2. For legend see facing page.

 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots 23

100

80

% of initial weight
60

40

20

0 20 40 60 80 100 120 140

Time (min)
F. 3. Water loss kinetics in basal (100–140 mm from roof apex) root segments from non-exodermal ($,
, >) and exodermal (#, , =) roots.

100 Zea mays

80
% of initial weight

60

40

20

0 20 40 60 80 100 120
Time (min)
F. 4. Water loss from apical (
) and basal ( ) segments of maize primary roots. Segments, 30 mm long, comprised the region located between
10–40 mm (apical) and 100–140 mm (basal) from the root apex. Results are meansps.e. of six measurements. Basal segments are from Fig. 3.

T     1. AŠerage weight retention as percentage of initial could reflect the contribution of the ‘ transendodermal
weight in serial measurements taken between 30 and 120 min, compartment ’ to water loss.
in apical and basal segments of pea roots There is controversial evidence on the water permeability
of the exodermis, the basis for the suggested protective role
Weight for this layer. Tests with isolated sleeves of epidermal\
Segment retention (%) t d.f. P hypodermal cells of maize (Clarkson et al., 1987), suggest
that the suberized walls remain relatively permeable, but
Apical 21n67p2n83 become impermeable if the roots are exposed to moist air
Basal 37n33p3n08 10n45 47 0n001
rather than being kept completely immersed in culture
t-test results are for paired apical and basal segments of six roots.
solution. Hydraulic conductivity measured in both hy-
podermal sleeves and in intact root segments depended on
the direction of flow and was lower in the outflow direction
(Shone and Clarkson, 1988). The correlation between the

F. 2. A, Cross-section of a Vicia faba root stained with berberine-toluidine blue and observed under UV illumination. Arrow indicates Casparian
band in the endodermis. Notice the absence of fluorescence in the hypodermis. B, Cross-section of an onion root stained as in A, arrow indicates
the exodermis. C, Cross-section of a control C. gayana root, stained as in A, arrow indicates the exodermis. D, As for C, but from a salinized
plant. E and F, Autofluorescence in cross-sections of roots from control (E) and droughted (F) sorghum plants. G and H, External view (G) and
cross-section (H) of a C. gayana root growing in dry soil. Observe the rhizosheath (r), aerenchyma (a) and epidermis-exodermis (e). Root diameter
2 mm.
24 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots

y = –0.61266 + 30.712x – 1.4931x2 R2 = 0.999

% weight retention after 30 min

80

60

40

20

0 1 2 3 4
Diameter (mm)
F. 5. Weight retention percentage in 12 % agar cylinders after 30 min of air exposure. Each point is the average of three cylinders.

90
% weight retention

70

50

30

0.5 0.7 0.9 1.1 1.3 1.5

Segment diameter (mm)
F. 6. Weight retention after 30 min of air exposure in root segments obtained between 80–140 mm from the apex. Each point is from a single
segment.
, >,
, Exodermal roots ; , =, #, non-exodermal roots.

presence of suberin in the hypodermal walls and the
presence of a Casparian band (Perumalla et al., 1990),
suggested the suberized exodermis contributes to protecting
the root against water loss in substrates with low water
availability. Nevertheless, it has been shown that the endo-
dermis in young Zea mays roots is a major barrier to the
movement of ions but not of water (Peterson, Murrmann
and Steudle, 1993). Sanderson (1983) reported the de-
velopment of State III (suberized and thickened wall with
substances of phenolic origin) endodermal cells was
correlated with decreases in water uptake by barley
seedlings, however, in maize the presence of suberin lamellae
was not strictly associated with low permeability to water or
solutes (Clarkson et al., 1987). Our results from water loss
measurements in exodermal and non-exodermal roots
support the suggested role of the exodermis in protecting
roots against water loss. In interpreting the results from
sorghum plants, the chemical nature of the exodermal
thickening should be considered, as thickness resulting from
increased lignin deposition would not constitute a high
resistance to the diffusion of uncharged molecules such as
water (Schreiber et al., 1994). Nevertheless, water retention
was higher in basal than in apical segments in both
exodermal and non-exodermal roots, indicating the presence
of the exodermis was not associated with this difference.
Instead, differences in tissue composition, cell maturity and
wall permeability may be responsible for the water retention
characteristics of those segments.
Tissue characteristics may also have influenced the
differences in rates of water loss observed between control
 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots 25

100

% of initial weight
80
*

60

0 20 40 60 80 100 120 140

Time (min)
F. 7. Water loss from basal segments of Chloris gayana nodal roots. Segments, 20 mm long, comprised the region located between 100–120 mm
from the root apex. Results are meansps.e. of eight–nine root segments obtained from C. gayana cv. Boma plants grown with the addition of
0 (Control) or 300 m NaCl (salt-treated), respectively. Asterisk indicates the first significant difference (P 0n05) between control and salt-
treated plants.

100

80
% of initial weight

60

40

20

0 20 40 60 80 100 120
Time (min)
F. 8. Water loss from basal segments of Sorghum bicolor nodal roots obtained from control and droughted plants. Segments, 20 mm long,
comprised the region located between 40–80 mm from the root apex. Results are meansps.e. of six segments.

and salinized roots, where faster water loss was expected in
roots of control plants, where the water potential gradient
with the medium was higher. The explanation may reside in
the lower degree of lacunar development observed in
salinized roots, as this tissue may constitute a barrier
against water loss (Moreshet, Huang and Huck, 1996).
The pathway for water movement is substantially
influenced by the driving force for water flow (Steudle,
1995). In the case of hydrostatic pressure gradients, it has
been suggested that water flow is through a predominantly
apoplasmic pathway ; in contrast, movement generated by
osmotic gradients seems to have a significant cell-to-cell
component (Steudle et al., 1993 ; Steudle, 1995 ; however, see
Jones et al., 1988). Water loss from roots is a diffusional
process, the observed rate difference in water loss between
exodermal and non-exodermal segments suggests, never-
theless, that the pathway for this process must also have a
relevant apoplastic component.
Rate of water loss is also a function of the diffusion
constant for the system (Cape and Percy, 1996), which is
affected by differences in specific area (area\mass) and by
differences in the diffusion rate per unit area. Diffusional
permeability, in turn, depends on the porosity of the tissue
and on the contribution of viscous flow through pores
(Shone and Clarkson, 1988). For basal segments, differences
in water retention associated with the presence of an
26 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots
exodermis were more significant than those associated with
segment diameter, implying that the exodermis has a role in
modifying water loss from roots.
Correlation between the presence of an exodermis and
plant habitat might be expected given the protective function
ascribed to this layer. However, Kroemer (1903, cited by
Perumalla et al., 1990) reported no correlation between
suberization of the exodermis and plant habitat. This raises
the question of the relevance of the exodermis in protecting
roots growing under field conditions against water loss.
Water exchange between the soil and roots is controlled by
the water potential and hydraulic conductance of three
components of the root-soil pathway : the root ; an air gap
that can form between roots and soil when roots growing in
drying soil shrink ; and the soil (Nobel and North, 1993).
The relative importance of these components varies with
soil hydration. Rhizosheaths form around the roots of
many plants growing in soil (McCully, 1995), particularly
under dry conditions (Nobel and North, 1993). Rhizo-
sheaths in Ferocactus acanthodes and Opuntia ficus-indica
play a more important role than the suberized endodermis
in restricting water loss (North and Nobel, 1992). In C.
gayana growing in very dry soil, roots were tightly covered
by soil sheaths. Sections of roots within soil sheaths were
moist and appeared healthy. In maize, the sheathed zone of
roots is associated with immature late metaxylem, and is
relatively isolated from the transpiration stream (Wang,
Canny and McCully, 1991) ; whether immature late
metaxylem also occurs in sheathed roots of C. gayana must
be investigated. Rhizosheaths, along with cortical lacunae
may act as the main barriers against water loss in these
roots. The exodermis with its engrossed walls could provide
additional reinforcement against radial collapse under dry
conditions and prevent the formation of air gaps in drying
soils.
A C K N O W L E D G E M E N TS
Support from Fundacio! n Antorchas is gratefully
acknowledged. L. Garcias efficiently helped at various
stages.
LITERATURE CITED
Baker JM, van Bavel CHM. 1986. Resistance of plant roots to water
loss. Agronomy Journal 78 : 641–644.
Blum A, Johnson B. 1992. Transfer of water from roots into dry soil and
the effect of wheat water relations and growth. Plant and Soil 145 :
141–149.
Boyer JS. 1985. Water transport. Annual ReŠiew of Plant Physiology
36 : 473–516.
Brundrett MC, Enstone DE, Peterson CA. 1988. A berberine-aniline
blue fluorescent staining procedure for suberin, lignin, and callose
in plant tissue. Protoplasma 146 : 133–142.
Cape JN, Percy KE. 1996. The interpretation of leaf-drying curves.
Plant, Cell and EnŠironment 19 : 356–361.
Clarkson DT, Robards AW, Stephens JE, Stark M. 1987. Suberin
lamellae in the hypodermis of maize (Zea mays) roots : development
and factors affecting the permeability of hypodermal layers. Plant,
Cell and EnŠironment 10 : 83–93.
Cruz RT, Jordan WR, Drew MC. 1992. Structural changes and
associated reduction of hydraulic conductance in roots of Sorghum
bicolor following exposure to water deficit. Plant Physiology 99 :
203–212.
Enstone DE, Peterson CA. 1992. The apoplastic permeability of root
apices. Canadian Journal of Botany 70 : 1502–1512.
Fro$ hlich MW. 1984. Freehand sectioning with Parafilm. Stain Tech-
nology 59 : 61–62.
Henberg HV. 1968. Der PrimaW rebau der Angiospermenwurzel. Berlin :
Gerbru$ der Borntra$ ger.
Jones H, Leigh RA, Wynn Jones RG, Tomos AD. 1988. The integration
of whole-root and cellular hydraulic conductivities in cereal roots.
Planta 174 : 1–7.
Jupp AP, Newman EI. 1987. Morphological and anatomical effects of
severe drought on the roots of Lolium perenne L. New Phytologist :
393–402.
Kikuta SB, Richter H. 1992. Leaf discs or press saps ? a comparison of
techniques for the determination of osmotic potentials in freeze-
thawed leaf material. Journal of Experimental Botany 43 :
1039–1044.
McCully M. 1995. How do real roots work ? Plant Physiology 109 : 1–6.
Melchoir W, Steudle E. 1993. Water transport in onion (Allium cepa
L.) roots. Changes of axial and radial hydraulic conductivities
during root development. Plant Physiology 101 : 1305–1315.
Moreshet S, Huang B, Huck MG. 1996. Water permeability in roots.
In : Waisel Y, Eshel A, Kafkafi U, eds. Plant roots : the hidden half.
New York : Marcel Dekker, 659–678.
Munns R, Passioura JB. 1984. Hydraulic resistance of plants. III.
Effects of NaCl in barley and lupin. Australian Journal of Plant
Physiology 11 : 351–359.
Nobel PS, North GB. 1993. Rectifier-like behaviour of root-soil
systems : new insights from desert succulents. In : Smith JAC,
Griffiths H, eds. Water deficits. Plant responses from cell to
community. Oxford : Bios Scientific Publishers, 163–186.
North GB, Nobel PS. 1992. Drought-induced changes in hydraulic
conductivity and structure in roots of Ferocactus acanthodes and
Opuntia ficus-indica. New Phytologist 120 : 9–19.
Oertli JJ. 1996. Transport of water in the rhizosphere and in roots. In :
Waisel Y, Eshel A, Kafkafi U, eds. Plant roots : the hidden half.
New York : Marcel Dekker, Inc., 607–633.
O’Brien TP, Feder N, McCully ME. 1964. Polychromatic staining of
plant cell walls by Toluidine Blue O. Protoplasma 59 : 369–373.
Passioura JB. 1988. Water transport in and to roots. Annual ReŠiew
Plant Physiology and Molecular Plant Biology 39 : 245–265.
Perumalla CJ, Peterson CA, Enstone DE. 1990. A survey of angiosperm
species to detect hypodermal casparian bands. I. Roots with a
uniseriate hypodermis and epidermis. Botanical Journal of the
Linnean Society 103 : 93–112.
Peterson CA. 1989. Significance of the exodermis in root function. In :
Loughman BC, ed. Structural and functional aspects of transport in
roots. Dordrecht : Kluwer Academic Publishers, 35–40.
Peterson CA, Enstone DE. 1996. Functions of passage cells in the
endodermis and exodermis roots. Physiologia Plantarum 97 :
592–598.
Peterson CA, Murrmann M, Steudle E. 1993. Location of the major
barriers to water and ion movement in young roots of Zea mays
L. Planta 190 : 127–136.
Peyrano G, Taleisnik E, Quiroga M, Forchetti SM, Tigier H. 1997.
Salinity effects on hydraulic conductance, lignin content and
peroxidase activity in tomato roots. Plant Physiology and
Biochemistry 35 : 387–393.
Reinhardt DH, Rost TL. 1995. Salinity accelerates endodermal
development and induces an exodermis in cotton seedling roots.
EnŠironmental and Experimental Botany 35 : 563–574.
Sanderson J. 1983. Water uptake by different regions of the barley root
in relation to development of the endodermis. Journal of
Experimental Botany 34 : 240–253.
Schreiber L, Breiner H-W, Riederer M, Du$ ggelin M, Guggenheim R.
1994. The Casparian strip of CliŠia miniata Reg. roots : isolation,
fine structure and chemical nature. Botanica Acta 107 : 353–361.
Shone MGT, Clarkson DT. 1988. Rectification of radial water flow in
the hypodermis of nodal roots of Zea mays. Plant and Soil 111 :
223–229.
 Taleisnik et al.—Water Loss in Exodermal and Non-exodermal Roots 27
Steudle E. 1992. The biophysics of plant water : compartmentation, solutes across maize roots modified by puncturing the endodermis.
coupling with metabolic processes, and flow of water in plant Plant Physiology 103 : 335–349.
roots. In : Somero G, Osmond CB, Bolis CL, eds. Water and life : Vetterlein D, Marschner H. 1993. Use of a microtensiometer technique
comparatiŠe analysis of water relationships at the organismic, to study hydraulic lift in a sandy soil planted with pearl millet
cellular and molecular leŠels. Heidelberg : Springer-Verlag, 173–204. (Pennisetum americanum). Plant and Soil 149 : 275–282.
Steudle E. 1995. Water transport across roots. Plant and Soil 167 : Wang X-L, Canny MJ, McCully ME. 1991. The water status of the
79–90. roots of soil-grown maize in relation to the maturity of their
Steudle E, Murrmann M, Peterson CA. 1993. Transport of water and xylem. Physiologia Plantarum 82 : 167–162.