Trees (2001) 15:335–340
DOI 10.1007/s004680100111
O R I G I N A L A RT I C L E
Osbert J. Sun · Gerty J. H. P. Gielen · Roger Sands
C. Tattersall Smith · Alan J. Thorn
Growth, Mg nutrition and photosynthetic activity in Pinus radiata:
evidence that NaCl addition counteracts the impact of low Mg supply
Received: 3 August 2000 / Accepted: 4 June 2001 / Published online: 10 July 2001
© Springer-Verlag 2001
Abstract We investigated effects of low Mg and moder-
ately raised NaCl, as occurs in plantations irrigated with
tertiary municipal effluent in New Zealand, on growth,
Mg nutrition and photosynthetic activity of Pinus radiata
D. Don seedlings grown in nutrient solutions with a
Perlite medium. Seedlings were grown with either suffi-
cient (0.35 mM; H[Mg]) or limited (0.033 mM; L[Mg])
Mg supply, without NaCl or with NaCl addition
(8.7 mM; +[NaCl]). After 30 weeks, seedlings grown at
L[Mg] displayed severe Mg deficiency symptoms, and
had significantly less biomass than those at H[Mg].
While NaCl addition had an adverse effect on seedling
growth at H[Mg], it increased growth at L[Mg]. The
+[NaCl] treatment greatly increased the Mg uptake rates,
which were associated with increased stomatal conduc-
tance and increased root to shoot ratio. Magnesium
deficiency reduced the rates of light-saturated photo-
synthesis and stomatal conductance, but not the quantum
efficiency of photosystem II, which was reduced mainly
by the +[NaCl] treatment, especially at H[Mg]. Our
study clearly indicated that NaCl addition could counteract
the impact of low Mg supply by enhancing Mg uptake
from the rooting medium.
Keywords Magnesium · Nutrition · Pinus radiata ·
Sodium · Solution culture
O.J. Sun (✉) · R. Sands
School of Forestry, University of Canterbury, Private Bag 4800,
Christchurch, New Zealand
e-mail: osbert.sun@orst.edu
Tel.: +1-541-7378471, Fax: +1-541-7371393
G.J.H.P. Gielen · A.J. Thorn
New Zealand Forest Research Institute Limited,
Private Bag 3020, Rotorua, New Zealand
C.T. Smith
Department of Forest Science, Texas A&M University,
College Station, TX 77843-2135, USA
Present address:
O.J. Sun, Department of Forest Science, Oregon State University,
Corvallis, OR 97331-5752, USA
Introduction
In New Zealand, there has been a large effort to utilize
forest plantations as a biofilter for treatment of domestic
and industrial liquid and solid wastes, to minimize the
impacts of these social and economic by-products on the
environment. Such schemes could provide a novel
system for removing and recycling pollutants using
non-food chain ecosystems, providing that any adverse
impacts of waste application on crop trees are carefully
considered. One concern is the impact on tree health and
nutrition resulting from modification of soil nutrient
supply. The problem could arise from either insufficient
supply of essential nutrients for growth, or from excessive
supply of toxic chemicals contained in the wastes.
Results from Pinus radiata D. Don plantations irrigated
with the tertiary municipal effluent from the sewage
treatment plant of Rotorua, New Zealand, showed
reduced foliar Mg concentrations, but significantly
increased soil and foliar Na concentrations (Gielen et al.
2000; McLay et al. 2000; Thorn et al. 2000) accompanied
also by increased Cl concentrations in soil solution
(Gielen et al. 2000). These observations raised issues
about the interactive effects of low Mg and raised NaCl
on the physiological performance and nutrition of
P. radiata, a predominant wood and fiber source in
New Zealand.
Magnesium plays an essential role in photosynthesis
and many other metabolic processes. In addition to being
the central component of the chlorophyll molecule, the
nutrient Mg is essential in enzymatic reactions in photo-
synthesis and in cellular pH control (Mengel and Kirby
1987; Marschner 1995). Magnesium deficiency has been
considered as the major factor in the wide occurrence of
needle yellowing and dieback of the upper-mid crown in
P. radiata stands in New Zealand (Beets and Jokela
1994). Magnesium deficiency causes impairment of
photosynthesis in P. radiata trees (Sun and Payn 1999;
Laing et al. 2000). Sodium is often associated with salinity
(Foster and Sands 1977; Sands and Clarke 1977), and is
toxic to many terrestrial plants at high concentrations. It
336
plays an important role in mediating membrane permeabi-
lity in plants for water and nutrient uptake, in contrast to
maintenance of the membrane integrity by Ca ion.
However, the requirements of most terrestrial plants for
Na are thought to be at the scale of trace levels (Mengel
and Kirby 1987).
Information is lacking on the interactive effects of Mg
and Na, and on the role of NaCl salinity in influencing
Mg uptake. Saur et al. (1995) found that high levels of
NaCl reduced Mg concentrations in roots of Pinus pinaster
Ait. seedlings. However, there is evidence that salinity
increases Mg uptake in some plants (Shukla and Singh
1996; Dahdoh and Hassan 1997).
To elucidate the implications of low Mg and raised
NaCl for growth and health of P. radiata plantations,
we studied growth and nutrition of seedlings grown at
sufficient or limited Mg supply, with or without NaCl
addition, in nutrient solutions with a Perlite medium.
The primary objective of our study was to investigate the
impacts of NaCl addition on Mg uptake and seedling
growth on sites with limited Mg supply. Although in
previous studies we have shown the effects of Mg
deficiency on P. radiata seedlings (Sun and Payn 1999;
Laing et al. 2000), it is not known that how NaCl addition
may further influence the impacts of Mg deficiency on
photosynthetic activities. Our second objective, therefore,
was to examine the interactive effects of Mg and NaCl
on the rates of CO2 fixation and quantum efficiency in
photosynthesis as a way of detecting any subtle changes
in physiological processes brought about by the imbalance
between the two chemicals.
Materials and methods
Plant culture and treatments
P. radiata seedlings were raised from control-pollinated seed
(GF28) in seed trays filled with potting mix. Two weeks after
emergence, they were transplanted into 4-l pots filled with Perlite
with two seedlings planted in each pot. The problem of water loss
from pots by evaporation was avoided by covering Perlite surface
with a 10-mm layer of polythene beads.
The experiment was conducted in a greenhouse between late
summer and early spring of the following year. Temperature in the
greenhouse was thermostatically controlled, but fluctuated
between 20°C and 25°C during the day, and 12°C and 15°C at
night depending on the weather conditions outside. Photoperiod
was extended to 16 h with 400-W sodium lamps.
The treatments were Mg [supplied as Mg(NO3)2; Sun and Payn
1999] concentrations at 0.35 mM (H[Mg]) and 0.033 mM
(L[Mg]), and without (–[NaCl]) or with NaCl addition at 8.7 mM
(+[NaCl]) in the cultural solutions, in a factorial arrangement with
five replications. The two levels of Mg concentration were chosen
to represent sufficient (0.35 mM) and limited (0.033 mM) levels
known to affect growth and physiological activities in P. radiata
(Sun and Payn 1999; Laing et al. 2000). Sodium chloride concen-
tration was chosen based on that typically found in the tertiary
municipal effluent (8.7 mM) from the sewage treatment plant of
Rotorua, New Zealand. Both Mg and NaCl were adjusted to the
specified concentrations in a base nutrient solution containing all
the other essential elements which were held constant relative to N
as defined by Ingestad (1971). The chemical composition of the
nutrient solutions was modified for the purpose of this study from
that reported by Sun and Payn (1999). Nitrogen concentrations in
all the treatment were maintained at 7.14 mM by varying the
amount of NH4NO3. The pH of the four treatment solutions ranged
from 5.3 to 5.5.
In the first 5 weeks following transplantation into 4-l pots, all
seedlings received a pre-treatment with a low strength
(3.6 mM[N]) of Ingestad’s (1971) complete nutrient solution, to
avoid any initial shock caused by full strength nutrient solutions.
Treatments commenced in the 6th week, initially with half
strength solutions for 4 weeks, followed by application of full
strength solutions for 15 weeks, on a weekly schedule with each
pot watered to the free-draining water-holding capacity of the
Perlite. For the 6 weeks prior to the harvest, treatment applications
were increased to twice a week to maintain a relatively stable
water and nutrient supply as demand increased with tree growth.
Growth and plant nutrient analysis
Seedlings were harvested after 30 weeks for measurement of
biomass and tissue nutrient analysis, on three subdivided tissue
components including needles, stem plus branches, and roots.
Tissues were analyzed for Mg, Na, N, P, K, Ca, B, Mn, Zn and Cu
by using the standard methods of Nicholson (1984). Uptake rates
of Mg (U[Mg]) per unit root dry weight (DWRt; g) were calculated
by the modified formula of Ingestad and Ågren (1988):
(1)
where C[Mg] is the integrated Mg concentration (mM) in seedlings,
accounting for the mass and Mg concentrations in needles, stem,
and roots, at harvesting; DWSl is the total seedling dry weight (g)
at harvesting; e the base of natural logarithm; and RGR is mean
relative growth rate, which is calculated as:
(2)
where DW0 is the initial total seedling dry weight (g), estimated
by destructively sampling six seedlings of similar size at the start
of the treatment; and ∆T is the duration of the experiment (days).
The calculations of both U[Mg] and RGR were made with the
assumption that the growth and nutrient uptake of seedlings were
at “steady state” during the entire period of experiment, considering
the relatively controlled nutrient application regimes and growth
environment within the greenhouse.
Photosynthesis and chlorophyll fluorescence
Measurements of photosynthetic gas exchange and chlorophyll
fluorescence were made in the greenhouse 1 week before seedlings
were harvested.
Rates of light-saturated net photosynthesis (A) and stomatal
conductance for diffusion of water vapor (gsw) were measured in
six fully expanded needles (two fascicles) at ambient CO2 concen-
tration in the greenhouse with a portable photosynthesis system
(LI-6400, Li-Cor, Lincoln, Neb., USA), and were expressed as per
unit surface needle area. The total surface area of the needle
segments enclosed in the leaf cuvette was determined with the
method described by Beets (1977). During measurements, the
photosynthetically active radiation (PAR) on the upper needle
surface was maintained at 1,000 µmol m–2 s–1 (the maximum level
experienced by seedlings in the greenhouse) using a red LED light
source (model LI–6400-02, Li-Cor, Lincoln, Neb., USA) attached
to the leaf cuvette, and temperature in the cuvette was set for 20°C.
Chlorophyll a fluorescence was measured in vivo with a portable
chlorophyll fluorometer (PEA Plant Efficiency Analyzer, Hansatech
Instruments, Kings Lynn, UK). Six fully expanded needles (two
fascicles) for each seedling were dark-adapted for more than
15 min. These needles were measured for the maximum and
minimum chlorophyll a fluorescence emissions (Fm and Fo) and
 337

Fig. 1 Dry weights of needles, stem, and roots at 30 weeks in

glasshouse-grown Pinus radiata D. Don seedlings at different
rates of Mg and NaCl supply. Vertical bars indicate standard
errors (n=5)

Table 1 Root to shoot ratio (R/S) of Pinus radiata D. Don seedlings

at 30 weeks as affected by Mg and NaCl supply. Values are means
(±SE; n=5)

R/S ratio % Increase

–[NaCl] +[NaCl]

H[Mg] 0.361 (±0.014) 0.410 (±0.013) 13.6%

L[Mg] 0.331 (±0.018) 0.420 (0.0±13) 26.9%

Fig. 2 Tissue Mg and Na concentrations in glasshouse-grown

Pinus radiata D. Don seedlings at different rates of Mg and NaCl
consequently the fluorescence yield (Fv=Fm – Fo) and the potential supplies. Vertical bars indicate standard errors (n=5)
maximal quantum efficiency (Fv/Fm) of photosystem II (PSII,
Schreiber et al. 1994). Fm was measured at a saturating irradiance
of 1,250 µmol m–2 s–1 provided by a built-in red LED light source. Table 2 Mean Mg uptake rate (U[Mg]) in Pinus radiata D. Don
seedlings as affected by Mg and NaCl supply. Values are means
(±SE; n=5)
Data analysis
U[Mg] (µM g–1DWroot day–1) % Increase
Data were evaluated by analysis of variance (ANOVA) for a
completely randomized design in a factorial arrangement with two –[NaCl] +[NaCl]
Mg concentrations, two NaCl concentrations and five replications.
The statistical analysis was performed with the ANOVA procedure H[Mg] 3.531 (±0.179) 4.853 (±0.190) 37.4%
of the SAS System for Windows release 6.12 (SAS 1989). L[Mg] 1.078 (±0.067) 1.475 (±0.056) 36.8%
% Decrease –69.5% –69.6%

Results
Biomass
Seedlings were less than 10 cm tall when treatments
were initialized. The rate of Mg supply, [Mg], had a
highly significant (P≤0.0001) effect on seedling biomass.
At 30 weeks, the dry weights of needles, stem and roots
of seedlings subjected to L[Mg] treatment were much
less than those at H[Mg] (Fig. 1). The reduction of biomass
in response to low Mg supply was associated with severe
Mg deficiency symptoms of typically yellowing needle
tips. The effect of NaCl addition, however, interacted
significantly (P≤0.05) with [Mg] for stem and roots; at
H[Mg], seedlings grown at +[NaCl] had lower dry
weights of needles, stem and roots, compared with those
grown at –[NaCl]; whilst at L[Mg], the dry weights of
stem and roots were 38 and 49% greater in seedlings
grown at +[NaCl] than at –[NaCl].
NaCl addition resulted in a highly significant
(P≤0.0001) increase in root to shoot ratio (R/S). The
increase was more pronounced at L[Mg] than at H[Mg]
(Table 1).
Tissue Mg and Na concentrations, and Mg uptake rates
Reducing [Mg] resulted in a substantial reduction
(P≤0.0001) in Mg concentration in all the three seedling
components, with the reduction being the most
pronounced in roots (Fig. 2). NaCl addition resulted in a
highly significant (P≤0.0001) increase in Mg concentration
of needles and roots, and a significant (P≤0.05) increase in
stem Mg concentration. The percentage of increase in Mg
concentration in response to the +[NaCl] treatment varied
among the three components, and was affected by [Mg].
Mg concentration was most responsive to NaCl addition
338
Table 3 Tissue concentrations of essential nutrients other than Mg and Na in needles, stem, and roots of Pinus radiata D. Don seedlings
grown at different rates of Mg and NaCl supply. Values are means (±SE; n=5)

Tissue Treatment Macro-nutrient (mg g–1 DW) Micro-nutrient (µg g–1 DW)

N P K Ca B Mn Zn Cu

Needle H[Mg]–[NaCl] 27.9 (±0.8) 3.05 (±0.08) 14.9 (±0.3) 1.25 (±0.04) 38.6 (±2.2) 140 (±6) 32.2 (±2.2) 20.8 (±2.2)
H[Mg]+[NaCl] 23.6 (±0.3) 2.60 (±0.06) 19.0 (±0.2) 1.67 (±0.10) 33.0 (±1.3) 156 (±6) 33.6 (±1.9) 16.0 (±1.7)
L[Mg]–[NaCl] 24.8 (±0.5) 3.30 (±0.04) 16.6 (±0.3) 1.31 (±0.08) 45.8 (±2.5) 211 (±20) 41.2 (±4.2) 57.9 (±1.8)
L[Mg]+[NaCl] 21.9 (±0.4) 2.86 (±0.04) 20.7 (±0.8) 2.54 (±0.11) 45 (±1.2) 234 (±6) 41.8 (±2.2) 21.6 (±2.8)
Stem H[Mg]–[NaCl] 9.9 (±0.7) 2.41 (±0.14) 19.0 (±0.6) 1.08 (±0.07) 18.2 (±0.7) 101 (±5) 29.4 (±2.5) 15.7 (±1.4)
H[Mg]+[NaCl] 8.3 (±0.2) 2.04 (±0.12) 20.1 (±1.1) 0.73 (±0.02) 16.6 (±0.3) 100 (±6) 27.8 (±1.0) 5.2 (±0.5)
L[Mg]–[NaCl] 13.0 (±1.0) 3.24 (±0.20) 18.3 (±1.7) 1.60 (±0.07) 17.2 (±2.9) 168 (±31) 48.2 (±9.7) 14.1 (±2.1)
L[Mg]+[NaCl] 9.0 (±0.2) 2.52 (±0.06) 21.0 (±1.1) 1.17 (±0.02) 18.4 (±1.0) 176 (±12) 42.6 (±1.6) 8.0 (±0.8)
Root H[Mg]–[NaCl] 17.4 (±0.5) 4.38 (±0.22) 16.3 (±0.7) 0.77 (±0.02) 19.4 (±1.5) 47 (±6) 31.2 (±4.3) 106.3 (±16.6)
H[Mg]+[NaCl] 17.5 (±0.6) 2.97 (±0.11) 12.1 (±0.6) 1.77 (±0.05) 29.8 (±1.3) 146 (±13) 38.0 (±1.7) 57.9 (±14.5)
L[Mg]–[NaCl] 20.5 (±0.8) 6.04 (±0.34) 17.2 (±0.7) 0.83 (±0.01) 13.4 (±1.2) 87 (±11) 62.2 (±5.2) 124.7 (±27.3)
L[Mg]+[NaCl] 18.9 (±0.3) 3.04 (±0.06) 14.9 (±0.5) 2.56 (±0.04) 25 (±0.9) 265 (±17) 52.8 (±2.0) 71.6 (±11.1)

Table 4 Summary of P (probability) value of treatment effects of rate of Mg and NaCl addition on tissue concentration of other nutri-
ents. (NS not significant at P>0.5)

Tissue Source Macro-nutrient Micro-nutrient

N P K Ca B Mn Zn Cu

Needle [Mg] <0.0001 0.0002 0.0016 <0.0001 <0.0001 <0.0001 0.0034 NS

[NaCl] <0.0001 <0.0001 <0.0001 <0.0001 NS NS NS 0.20
[Mg]×[NaCl] NS NS NS <0.0001 NS NS NS NS
Stem [Mg] 0.0039 <0.0001 NS <0.0001 NS 0.0003 0.002 NS
[NaCl] 0.0002 0.0003 NS <0.0001 NS NS NS <0.0001
[Mg]×[NaCl] 0.0490 NS NS NS NS NS NS NS
Root [Mg] 0.0004 0.0003 0.0035 <0.0001 0.0002 <0.0001 <0.0001 NS
[NaCl] NS <0.0001 <0.0001 <0.0001 <0.0001 <0.0001 NS 0.0071
[Mg]×[NaCl] NS 0.0007 NS <0.0001 NS 0.0022 0.0237 NS

in roots, followed by needles, with the stem Mg concentra-
tion being the least susceptible (Fig. 2).
The rate of Mg uptake, U[Mg], was highly significantly
(P≤0.0001) affected by both [Mg] and [NaCl]. U[Mg]
decreased with decreasing [Mg], and increased by NaCl
addition (Table 2).
The rates of [Mg] and [NaCl] addition also affected
the concentrations of nutrient elements other than Mg
and Na in all tissues (Table 3). A summary of the statistical
analysis of the treatment effects on eight essential
macro- and micronutrients is given in Table 4.
Photosynthesis and quantum efficiency
The rate of net photosynthesis, A, decreased significantly
(P≤0.001) with decreasing [Mg] (Fig. 3). A was also
affected (P=0.053) by an interaction between [Mg] and
[NaCl]; at H[Mg], [+NaCl] reduced A by an average of
20%; whereas at L[Mg], A was increased by an average
of 18% with +[NaCl]. The stomatal conductance to
diffusion of water vapor, gsw, decreased significantly
(P≤0.001) with decreasing [Mg], but increased by
+[NaCl].
Fv/Fm was affected by [NaCl] (P≤0.0005), and by
an interaction between [Mg] and [NaCl] (P≤0.05).
Fv/Fm decreased slightly with decreasing [Mg] at
–[NaCl], but increased with decreasing [Mg] at +[NaCl]
(Fig. 3).
Discussion
The impact of imbalanced nutrient loading on tree
growth and health is a concern with land application of
wastes to forest plantations. The potential risks include
deficiency of essential nutrients and toxicity of undesirable
chemicals, such as the conditions of low Mg and high
NaCl supply.
Magnesium deficiency and NaCl toxicity are both
known to affect plant growth. However, the interactive
effects of the two conditions on growth and physiological
activities in trees are poorly understood. In this study, we
found that although the growth of P. radiata seedlings
was inhibited by low Mg supply, and by NaCl addition
when Mg supply was not limited, the effect of low Mg
supply on growth could be partly counteracted by NaCl
addition. Under deficient Mg supply, raising NaCl supply

Fig. 3 Light-saturated rates of net photosynthesis (A), stomatal
conductance to diffusion of water vapor (gsw), and quantum
efficiency of PSII (Fv/Fm) in glasshouse-grown Pinus radiata
D. Don seedlings at different rates of Mg and NaCl supply. Vertical
bars indicate standard errors (n=5)
increased stem and root biomass, and increased Mg
uptake rates, by ≈40%.
Sodium is known to play a key role in mediating the
permeability of plasma membranes, in contrast to mainte-
nance of the membrane integrity by Ca. The enhance-
ment of Mg uptake by NaCl addition in P. radiata found
in our study may result from the effect of Na on mem-
brane permeability of roots. This differs from previously
documented studies of interactions between Mg and
other cations (Mengel and Kirby 1987; Puech and
Mehne-Jacobs 1997; Sun and Payn 1999). The uptake of
Mg in roots is generally slow compared with uptake of
other nutrients due to lack of a specific uptake and trans-
port mechanism across plasma membrane (Mengel and
Kirby 1987). The process is considered to be passive and
operate through electrochemical gradients across the ion
pores in the membranes, although an active uptake
component was suggested to exist by a study using radio-
active isotope 24Mg (Hogberg et al. 1995). The uptake of
Mg has been found to be complex and relatively slow in
Pinus taeda L. seedlings (Kelly and Barber 1991; Kelly
et al. 1992). Potassium has been shown to interfere with
both uptake and mobilization of Mg in tissue-cultured
P. radiata clones (Sun and Payn 1999). In the present
study, however, the addition of another monovalent
cation, Na, enhanced the uptake of Mg. Although there
is no direct evidence on increased permeability of Mg
339
across the plasma membrane by Na, alteration of the ion
transport systems of the cell plasmalemma by other
cations has been found to exist in plants. It was found
that, in a freshwater alga (Nitella flexilis), excessive Cu
increased plasmalemma conductance that was nonselective
and potential independent (Demidchik et al. 1997). The
increased stomatal conductance found in this study may
indirectly indicate increased membrane permeability in
roots of P. radiata seedlings grown with moderate NaCl
addition.
The potential effect of Cl on Mg nutrition in P. radiata
was not easily identifiable in the present study. Foster and
Sands (1977) suggested that P. radiata was adapted to
avoid or delay the accumulation of Cl in vital tissues, partly
by preventing Cl from entering the roots. However, we
lacked data on tissue Cl concentrations in both field-
grown P. radiata trees irrigated with tertiary municipal
effluent and in our greenhouse study for an explicit inter-
pretation of the role of Cl in mediating Mg uptake. Further
studies are needed to prove that the effects we observed
from NaCl addition were from Na and not Cl. However,
some studies showed that Na and Cl have different effects
on growth and nutrition in plants (Marschner 1995; Martin
and Koebner 1995), with the former element having a
positive effect on uptake of other nutrients.
The markedly reduced seedling growth with low Mg
supply apparently results from the decline in photo-
synthetic CO2 fixation rate (Sun and Payn 1999; Laing et
al. 2000). The Mg-deficiency-induced reduction of
photosynthesis in conifers has been suggested to result
from blockage of carbohydrate exportation from affected
leaves (Mehne-Jakobs 1995, 1996), possibly due to
damage to vascular tissues (Hannick et al. 1993). Similar
to the study of Mehne-Jakobs (1996) with Picea abies
(L.) Karst., we found Mg deficiency did not affect
quantum efficiency of PSII of Pinus radiata. Results
also showed that the reduction of photosynthesis in
response to low Mg supply was associated with reduction
of stomatal conductance, in agreement with results from
previous studies (Sun and Payn 1999; Laing et al. 2000).
Sodium is considered to be a beneficial mineral element
to certain plants (Marschner 1995), but has often been
associated with salinity stress at high concentrations
(Halperin et al. 1997; Sands and Clarke 1977). In green-
house-grown P. radiata seedlings, the effect of NaCl on
photosynthesis and growth interacted with the level of
Mg supply. At sufficient Mg supply, NaCl addition
resulted in markedly reduced net photosynthetic rate,
quantum efficiency of PSII, and biomass production;
whereas at low Mg supply, NaCl addition increased the
rate of net photosynthesis, and biomass production of
stem and roots, although the quantum efficiency of PSII
was reduced.
We found that NaCl addition significantly reduced
quantum efficiency of PSII in P. radiata, and the effect
was greater when Mg supply was high. This significant
reduction of both light-saturated net photosynthesis and
the quantum efficiency of PSII by NaCl addition at high
Mg supply could result from excessive Mg.
340
Our study clearly indicated that NaCl addition could
counteract the effects of low Mg supply by enhancing
the uptake rate of the nutrient. We conclude that the level
of NaCl in the tertiary municipal effluent will not, at a
minimum, adversely impact on Mg nutrition of P. radiata
in a system where P. radiata plantations are used for the
treatment of such wastes.
Acknowledgements This research was funded by the Foundation
for Research, Science and Technology of New Zealand Contract
C04409. We thank Dr W. Laing and three anonymous reviewers
for constructive comments on an early version of the manuscript.
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