Escolar Documentos
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DOI 10.1007/s004680100111
O R I G I N A L A RT I C L E
Received: 3 August 2000 / Accepted: 4 June 2001 / Published online: 10 July 2001
© Springer-Verlag 2001
plays an important role in mediating membrane permeabi- that reported by Sun and Payn (1999). Nitrogen concentrations in
lity in plants for water and nutrient uptake, in contrast to all the treatment were maintained at 7.14 mM by varying the
amount of NH4NO3. The pH of the four treatment solutions ranged
maintenance of the membrane integrity by Ca ion. from 5.3 to 5.5.
However, the requirements of most terrestrial plants for In the first 5 weeks following transplantation into 4-l pots, all
Na are thought to be at the scale of trace levels (Mengel seedlings received a pre-treatment with a low strength
and Kirby 1987). (3.6 mM[N]) of Ingestad’s (1971) complete nutrient solution, to
avoid any initial shock caused by full strength nutrient solutions.
Information is lacking on the interactive effects of Mg Treatments commenced in the 6th week, initially with half
and Na, and on the role of NaCl salinity in influencing strength solutions for 4 weeks, followed by application of full
Mg uptake. Saur et al. (1995) found that high levels of strength solutions for 15 weeks, on a weekly schedule with each
NaCl reduced Mg concentrations in roots of Pinus pinaster pot watered to the free-draining water-holding capacity of the
Ait. seedlings. However, there is evidence that salinity Perlite. For the 6 weeks prior to the harvest, treatment applications
were increased to twice a week to maintain a relatively stable
increases Mg uptake in some plants (Shukla and Singh water and nutrient supply as demand increased with tree growth.
1996; Dahdoh and Hassan 1997).
To elucidate the implications of low Mg and raised
NaCl for growth and health of P. radiata plantations, Growth and plant nutrient analysis
we studied growth and nutrition of seedlings grown at
Seedlings were harvested after 30 weeks for measurement of
sufficient or limited Mg supply, with or without NaCl biomass and tissue nutrient analysis, on three subdivided tissue
addition, in nutrient solutions with a Perlite medium. components including needles, stem plus branches, and roots.
The primary objective of our study was to investigate the Tissues were analyzed for Mg, Na, N, P, K, Ca, B, Mn, Zn and Cu
impacts of NaCl addition on Mg uptake and seedling by using the standard methods of Nicholson (1984). Uptake rates
of Mg (U[Mg]) per unit root dry weight (DWRt; g) were calculated
growth on sites with limited Mg supply. Although in by the modified formula of Ingestad and Ågren (1988):
previous studies we have shown the effects of Mg
deficiency on P. radiata seedlings (Sun and Payn 1999; (1)
Laing et al. 2000), it is not known that how NaCl addition
may further influence the impacts of Mg deficiency on where C[Mg] is the integrated Mg concentration (mM) in seedlings,
photosynthetic activities. Our second objective, therefore, accounting for the mass and Mg concentrations in needles, stem,
was to examine the interactive effects of Mg and NaCl and roots, at harvesting; DWSl is the total seedling dry weight (g)
at harvesting; e the base of natural logarithm; and RGR is mean
on the rates of CO2 fixation and quantum efficiency in relative growth rate, which is calculated as:
photosynthesis as a way of detecting any subtle changes
in physiological processes brought about by the imbalance (2)
between the two chemicals.
where DW0 is the initial total seedling dry weight (g), estimated
by destructively sampling six seedlings of similar size at the start
Materials and methods 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
Plant culture and treatments at “steady state” during the entire period of experiment, considering
the relatively controlled nutrient application regimes and growth
P. radiata seedlings were raised from control-pollinated seed environment within the greenhouse.
(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 Photosynthesis and chlorophyll fluorescence
from pots by evaporation was avoided by covering Perlite surface
with a 10-mm layer of polythene beads. Measurements of photosynthetic gas exchange and chlorophyll
The experiment was conducted in a greenhouse between late fluorescence were made in the greenhouse 1 week before seedlings
summer and early spring of the following year. Temperature in the were harvested.
greenhouse was thermostatically controlled, but fluctuated Rates of light-saturated net photosynthesis (A) and stomatal
between 20°C and 25°C during the day, and 12°C and 15°C at conductance for diffusion of water vapor (gsw) were measured in
night depending on the weather conditions outside. Photoperiod six fully expanded needles (two fascicles) at ambient CO2 concen-
was extended to 16 h with 400-W sodium lamps. tration in the greenhouse with a portable photosynthesis system
The treatments were Mg [supplied as Mg(NO3)2; Sun and Payn (LI-6400, Li-Cor, Lincoln, Neb., USA), and were expressed as per
1999] concentrations at 0.35 mM (H[Mg]) and 0.033 mM unit surface needle area. The total surface area of the needle
(L[Mg]), and without (–[NaCl]) or with NaCl addition at 8.7 mM segments enclosed in the leaf cuvette was determined with the
(+[NaCl]) in the cultural solutions, in a factorial arrangement with method described by Beets (1977). During measurements, the
five replications. The two levels of Mg concentration were chosen photosynthetically active radiation (PAR) on the upper needle
to represent sufficient (0.35 mM) and limited (0.033 mM) levels surface was maintained at 1,000 µmol m–2 s–1 (the maximum level
known to affect growth and physiological activities in P. radiata experienced by seedlings in the greenhouse) using a red LED light
(Sun and Payn 1999; Laing et al. 2000). Sodium chloride concen- source (model LI–6400-02, Li-Cor, Lincoln, Neb., USA) attached
tration was chosen based on that typically found in the tertiary to the leaf cuvette, and temperature in the cuvette was set for 20°C.
municipal effluent (8.7 mM) from the sewage treatment plant of Chlorophyll a fluorescence was measured in vivo with a portable
Rotorua, New Zealand. Both Mg and NaCl were adjusted to the chlorophyll fluorometer (PEA Plant Efficiency Analyzer, Hansatech
specified concentrations in a base nutrient solution containing all Instruments, Kings Lynn, UK). Six fully expanded needles (two
the other essential elements which were held constant relative to N fascicles) for each seedling were dark-adapted for more than
as defined by Ingestad (1971). The chemical composition of the 15 min. These needles were measured for the maximum and
nutrient solutions was modified for the purpose of this study from minimum chlorophyll a fluorescence emissions (Fm and Fo) and
337
–[NaCl] +[NaCl]
Results
NaCl addition resulted in a highly significant
Biomass (P≤0.0001) increase in root to shoot ratio (R/S). The
increase was more pronounced at L[Mg] than at H[Mg]
Seedlings were less than 10 cm tall when treatments (Table 1).
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 Tissue Mg and Na concentrations, and Mg uptake rates
of seedlings subjected to L[Mg] treatment were much
less than those at H[Mg] (Fig. 1). The reduction of biomass Reducing [Mg] resulted in a substantial reduction
in response to low Mg supply was associated with severe (P≤0.0001) in Mg concentration in all the three seedling
Mg deficiency symptoms of typically yellowing needle components, with the reduction being the most
tips. The effect of NaCl addition, however, interacted pronounced in roots (Fig. 2). NaCl addition resulted in a
significantly (P≤0.05) with [Mg] for stem and roots; at highly significant (P≤0.0001) increase in Mg concentration
H[Mg], seedlings grown at +[NaCl] had lower dry of needles and roots, and a significant (P≤0.05) increase in
weights of needles, stem and roots, compared with those stem Mg concentration. The percentage of increase in Mg
grown at –[NaCl]; whilst at L[Mg], the dry weights of concentration in response to the +[NaCl] treatment varied
stem and roots were 38 and 49% greater in seedlings among the three components, and was affected by [Mg].
grown at +[NaCl] than at –[NaCl]. 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)
N P K Ca B Mn Zn Cu
in roots, followed by needles, with the stem Mg concentra- Fv/Fm was affected by [NaCl] (P≤0.0005), and by
tion being the least susceptible (Fig. 2). an interaction between [Mg] and [NaCl] (P≤0.05).
The rate of Mg uptake, U[Mg], was highly significantly Fv/Fm decreased slightly with decreasing [Mg] at
(P≤0.0001) affected by both [Mg] and [NaCl]. U[Mg] –[NaCl], but increased with decreasing [Mg] at +[NaCl]
decreased with decreasing [Mg], and increased by NaCl (Fig. 3).
addition (Table 2).
The rates of [Mg] and [NaCl] addition also affected
the concentrations of nutrient elements other than Mg Discussion
and Na in all tissues (Table 3). A summary of the statistical
analysis of the treatment effects on eight essential The impact of imbalanced nutrient loading on tree
macro- and micronutrients is given in Table 4. 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
Photosynthesis and quantum efficiency chemicals, such as the conditions of low Mg and high
NaCl supply.
The rate of net photosynthesis, A, decreased significantly Magnesium deficiency and NaCl toxicity are both
(P≤0.001) with decreasing [Mg] (Fig. 3). A was also known to affect plant growth. However, the interactive
affected (P=0.053) by an interaction between [Mg] and effects of the two conditions on growth and physiological
[NaCl]; at H[Mg], [+NaCl] reduced A by an average of activities in trees are poorly understood. In this study, we
20%; whereas at L[Mg], A was increased by an average found that although the growth of P. radiata seedlings
of 18% with +[NaCl]. The stomatal conductance to was inhibited by low Mg supply, and by NaCl addition
diffusion of water vapor, gsw, decreased significantly when Mg supply was not limited, the effect of low Mg
(P≤0.001) with decreasing [Mg], but increased by supply on growth could be partly counteracted by NaCl
+[NaCl]. addition. Under deficient Mg supply, raising NaCl supply
339
Our study clearly indicated that NaCl addition could Laing W, Greer D, Sun O, Beets P, Lowe A, Payn T (2000) Physio-
counteract the effects of low Mg supply by enhancing logical impacts of Mg deficiency in Pinus radiata: growth and
photosynthesis. New Phytol 146:47–57
the uptake rate of the nutrient. We conclude that the level McLay CDA, Tomer MD, Hopkins KM, Smith CT, Thorn AJ
of NaCl in the tertiary municipal effluent will not, at a (2000) Chemical changes in a volcanic forest soil after four
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in a system where P. radiata plantations are used for the Bastian RK (eds) The forest alternative: principles and practice
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