Functional
Ecology 2004
18, 419 – 425
© 2004 British
Ecological Society
419
Blackwell Publishing, Ltd.
Allocation of nitrogen to cell walls decreases
photosynthetic nitrogen-use efficiency
Y. ONODA,† K. HIKOSAKA and T. HIROSE
Graduate School of Life Sciences, Tohoku University, Aoba, Sendai, 980-8578, Japan
Summary
1. Nitrogen (N) is an essential limiting resource for plant growth, and its efficient use
may increase fitness. We investigated photosynthetic N-use efficiency (photosynthetic
capacity per unit N) in relation to N allocation to Rubisco and to cell walls in Polygonum
cuspidatum Sieb. et Zucc. which germinated in May (early germinators) and August
(late germinators).
2. There was a significant difference between early and late germinators in photo-
synthetic capacity as a function of leaf N content per unit area. Higher photosynthetic
N-use efficiency in late germinators was caused primarily by a larger allocation of N
to Rubisco.
3. Nitrogen allocation to cell walls was smaller in late germinators. The shorter growth
period in late germinators was associated with higher photosynthetic capacity, which
was achieved by allocating more N to photosynthetic proteins at the expense of cell
walls.
4. The trade-off between N allocation to photosynthesis and to structural tissues
suggests that plants change N allocation to increase either the rate or duration of
carbon assimilation. Such plastic change would help plants maintain themselves and
cope with environmental changes.
Key-words: cell-wall proteins, germination time, leaf life span, nitrogen allocation, ribulose-1,5-bisphosphate
carboxylase/oxygenase
Functional Ecology (2004) 18, 419– 425
Introduction
Nitrogen availability often limits plant growth in
natural ecosystems and its efficient use may increase
fitness (Chapin 1980; Aerts & Chapin 2000). Leaves
accumulate most of the N in the plant, and about half
the total leaf N is used for photosynthetic activities.
Thus the photosynthetic apparatus is the largest sink
of N in the plant (Evans & Seemann 1989; Poorter &
Evans 1998). Ribulose-1,5-bisphosphate carboxylase/
oxygenase (Rubisco) constitutes ≈ 50% of photosynthetic
N, and is the most abundant and important enzyme
of photosynthesis (Evans 1989). Consequently, photo-
synthetic capacity correlates strongly with leaf N con-
centration (Field & Mooney 1986; Evans 1989).
It is less clear how much leaf N is allocated to
components involved in activities other than photo-
synthesis. In addition to many nitrogenous compounds
required for cellular metabolism, cell walls are a major
N sink (Lambers & Poorter 1992). Cell walls contain
proteins that function in defence, growth, development,
signalling, intercellular communication and environ-
mental sensing, and as selective exchange interfaces
(Showalter 1993). Lamport (1965) showed that primary
cell walls have a substantial amount of N, ranging from
0·4–2·2% in several species. If cell walls account for
30–50% of leaf dry mass (Merino, Field & Mooney
1984), they constitute a considerable amount of leaf N.
Leaves with a long life span tend to have a high leaf
mass per area (LMA) and low photosynthetic N-use
efficiency (PNUE, photosynthetic capacity per unit
N). LMA, which is positively correlated with struc-
tural toughness, is often used as an index of structural
(cell-wall) biomass (Reich et al . 1991; Wright & Cannon
2001). On the other hand, LMA is negatively correlated
with PNUE (Poorter & Evans 1998). These studies sug-
gest that leaves with a long life span invest more N in
cell walls and consequently less in the photosynthetic
apparatus. Thus a trade-off between N allocation to
cell walls and to Rubisco is expected across leaves with
different life spans.
Leaf life span of a species changes with biotic and
abiotic factors. Germination time is one of the factors
that determine leaf life span in deciduous plants. Even
when they germinate at different times, they shed all
their leaves at the end of the growing season. Con-
sequently late germinators have a shorter leaf life span,
†Author to whom correspondence should be addressed.
E-mail: onoda@mail.tains.tohoku.ac.jp