Acta (Ecologica, 1997, 18 (2), 73-90

Nutrient dynamics during the development of a Douglas-fir

(Pseudotsuga menziesii Mirb.) stand

J. Ranger*, R. Marques and M. Colin-Belgrand

1NRA Centre de Nancy, Equipe "Cycles biog~ochimiques", 54280 Champenoux, France.
* Corresponding auth~

Received: 25.4.96 Accepted: 21.11.96

Abstract
The general process of nutrient cycling in forest stands is relatively well-known. Nevertheless,
fluxes in representative ecosystems have not yet been quantified, certainly due to the amount of mea-
surements required. The unavailability of such information is delaying the validation of models
describing the ecosystem function.
Numerous observations of nutrient stocks and fluxes carried out on a chronosequence of Dou-
glas-fir stands in the Beaujolais mountains (France) were used: i) to quantify the main fluxes of the
biological cycle of nutrients for the main stages of development of a single stand, ii) to characterize
their dynamics during forest development and iii) to quantify the real amount of nutrients involved in
forest growth and to identify their origins.
The present soil nutrient reserves of the ecosystem studied were relatively low if compared to
stand nutrient uptake. The biological cycle represents a key process in the maintenance of forest pro-
ductivity under such conditions. Cumulation of fluxes during rotation was very useful in quantifying
the real amounts of nutrients involved in tree nutrition, in demonstrating the efficiency of the system's
use of the limited pool o f nutrients needed to produce biomass and to show the relevance of the bio-
logical cycle in the soil function.
These results will be helpful in understanding the nutrient behaviour in a highly productive forest
plantation and thereby providing decisive information for their sustainable management.

Keywords: Biological cycle, nutrients, forest stand, requirements, uptake, internal translocation, lit-
terfall, Douglas-fir.

R~sum~
Le fonctiormement du cycle biologique des 616ments nutritifs dans les peuplements forestiers est
relativement bien connu. Cependant, la quantification des flux dans des ~cosyst~mes repr6sentatifs est
loin d'avoir 6t6 r6alis6e. Ce manque d'information est de nature h retarder la validation des modules
d6crivant le fonctionnement des 6cosyst~mes forestiers.
Les nombreuses observations concemant les stocks et les flux dans une chronos6quence de peu-
plements de Douglas darts les Monts du Beaujolais (France) ont 6t6 utilis6es : i) pour quantifier les
principaux flux du cycle biologique des 616ments nutritifs pour les principanx stades de d6veloppe-

Acta CEcologica 1146-609 X/97/02/$ 7.00 9 Gauthier-Villars

74 j . Ranger etal.

ment d'un m~me peuplement, ii) pour caractrriser la dynamique de ces 616ments au cours de la r6volu-
tion forestirre et iii) pour quantifier la quantit6 rrelle d'616ments raise en jeu dans la production
foresfi~re et en ~identlfier les origines.
Les rSserves en 616ments nutritifs des sols de l'rcosystSme &udi6 sont relativement faibles par
rapport au prrlbvement des peuplements. Le cycle biologique des 616ments reprrsente un processus-
c16 pour te maintien de la production dans ces conditions de fertilitr. Le cumul des flux durant la r~vo-
lution foresti~re a pennis i) de quantifier la masse rrelle d'616ments mise en jeu pour l'alimentation des
peuplemcnts, ii) de drrnontrer l'efficience du systSme pour utiliser le pool limit6 d'616ments n:utritifs
pour produire :la biomasse et iii) de mettre en 6vidence le rOle du cycle biologique dans le foncfionne-
ment des sols.
Ces rrsultats seront d'une grande utilit6 pour comprendre le fonctionnement mindral des planta-
tions forestiSres de forte production, apportant ainsi une information drterminante pour la gestion
durable de ces 6cosystSmes.

INTRODUCTION

The biological cycle of nutrients is a very efficient mechanism allowing tree

growth in soil with limited nutrient resources. Relatively high nutrient demand of
stands is satisfied for certain nutrients like N, P and K, both by uptake from soil and
by internal translocations within the plant itself, allowing a re-use of nutrients when
tissues cease to be physiologically active (HELMISAARI,1992). A large part of nutrients
taken up from the soil return to the forest floor as litterfall. This mechanism contrib-
utes to maintaining a satisfactory base cation saturation of the adsorbing complex in
the upper soil horizons of acidic soils (RANCER& COLIN-BELGRAND,1996).
Nutrient Cycling is relatively well known, especially after the research initiated
by the PBI/UNESCO reported in the synthesis of COLE & RAeP (1981), or obtained in
mid-term ecosystem study projects (DuVlCNEAUD,1975; LICKENSet al., 1977; I~ANNA
& ULI~CH, i991). Nevertheless, fluxes in representative ecosystems and their
dynamics during stand development have not yet been fully quantified, certainly due
to the amount of measurements necessary and the longevity of forest stands. The
unavailability of such information is delaying the validation of models describing the
ecosystem function.
Numerous observations of nutrient stocks and fluxes carried out on a chronose-
quence of Douglas=fir (Pseudotsuga menziesii Mirb.) stands in the Beaujolais moun-
tains (France), allowed us to quantify the main fluxes of the biological cycle of
nutrients for the main stages of development of a single stand. Nutrient requirements
for annual biomass production, uptake of vegetation from the pool of available soil
nutrients, internal translocation fi'om old to young tissues, return to the soil from lit-
terfall and crown leaching by rain were evaluated.
The objectives of this paper were i) to quantify the main nutrient fluxes of the
biological cycle in a Douglas-fir ecosystem, ii) to characterize their dynamics during
forest development, iii) to evaluate the real amount of nutrients involved in forest
growth and iv) to identify and quantify the different sources of these nutrients.
These results will be helpful in understanding nutrient behaviour in a highly pro-
ductive forest plantation and thus provide relevant information for the sustainable
management of these stands.

Acta (Ecologica
Nutrient dynamics in a forest stand 75

MATERIALS AND METHODS

Site and stands

The site studied is situated in the forest of Les Aiguillettes in the Beaujolais mountains (Rh6ne -
France). The elevation is about 750 m and the climate is characterised by a mean annual temperature of
7~ and a mean annual precipitation of 1000 ram.
The stands selected were 20, 40 and 60 years-old in 1990. They were all first rotations of Dou-
glas-fir planted after Spruce (20 and 40 year-old stands) or directly on land abandoned by agriculture
(60 year-old stand). The main characteristics of the stands are given in table I. The position of these
stands in the relationship between dominant height and age put them into fertility class 1 of the Dou-
glas-fir yield table established for the north-east Massif Central (France) by DECOURT(1967).

TABLEI. -- General stand characteristics.

stand age (years) 20 40 60
mean height (m) 14.3 28.0 36.0
mean Circumference at 1.3 m C13o (cm) 5710 104.7 163.7
stand basal area G (m2) 24.2 47.4 64.8
stand density (No of trees per ha) 922 490 312

The geological substratum was a Devono-Dinantian volcanic "tuff" relatively rich in base cations
(1.8% CaO, 1.8% MgO). The soil was of the Alocrisol type (AFES, 1992) with a loamy texture and a
rather high stone content (20 to 50% according to soil horizons and locations). The main soil character-
istics from the three stands were the following: the soil was acidic with a p H H 2 0 of the solid phase
ranging from 4.2 in the A horizon to 4.6 in the/3 horizon ; the base cation content was low from 8 to
1l tamale kg -1 for Ca and about 2 mmolc kg -1 for Mg in A1 and from 1.4 to 6.8 mmolc kg -1 f o r C a
and from 0.5 to 3 mmolc kg- 1 for Mg in B. Exchangeable AI occupied the largest part of the sites of the
cation exchange c~pacity (CEC) with values ranging from 35 to 70 mmolc kg -1 according to soil
layers.
Humus belonged to the mull/reader type with a total organic carbon content ranging from 40 to
85 mg g-1 and a C / N ratio of about 12.5 in the A l horizon.
The analysis of variance made on seven soil profiles individually analysed on each stand showed
a tendency to a more elevated base cation saturation on the soil of the young stand. This was attributed
to specific local colluvial deposits.
The available soil reserves for plants (i.e. exchangeable cations Ca, Mg and K), available phos-
phorus (DuclqAUFOuR-Bor,n~mAUmethodology, 1959) and the total amount of nitrogen are presented in
table II.

Methodology
The chronosequence approach has been used for a long time by forest research due to forest stand
longevity (COLE & VAN MmrROET, 1989). This is still the most suitable way to study the dynamic
aspect of forest ecosystem function, but it remains an approximation.
Thebiomass and nutrient content measurements were described in detail in RANCER et aL (1995).
The nutrient content of ~he stand was evaluated from destructive sampling: 12 trees per stand distrib-
uted throughout the girth classes defined from the inventory were cut down and the major cam-

Vol. I8, n o 2 - 1997

76 J. Ranger et aL

TABLEII. - Soil reserves in the organic layers and on the mineral soil (data in kg per ha for lm depth).

Dry malter N P K Ca Mg
20-year-old stand
L layer 4386 69 4 5 42 5
F+H layer (1) 21032 311 23 120 105 75
mineral soil (1m) 7700 1322 286 573 133
TOTAL 8080 1349 411 721 213
40-year-old stand
L layer 12395 149 8 21 48 8
F+H layer (1) 42221 524 35 195 90 90
mineral soil (lm) 5970 507 354 355 50
TOTAL 6642 550 569 493 147
60-year-old stand
L layer 7745 88 6 10 53 4
F+H layer (1) 47213 500 36 413 145 138
mineral soil (lm) 7480 370 594 614 132
TOTAL 8068 412 1017 812 275
(1) Total content of the layer including the mineral fraction representing approximately half of the mass.

ponents were isolated (needles one year old or more, branches one year old or more, stemwood and
stembark). Intensive measurements were taken on these trees: stem length, circumferences at different
levels on the stem, weight, branch diameter 10 cm from insertion on stem and position. Samples were
taken from all components for volume increment, wood and bark proportion and density (stem), needle
and wood proportion (branches), moisture and nutrient content (all). Common biomass and nutrient
content tables for the three stands were calculated. These tables were then applied to the inventory of
the stands in order to estimate their biomass and nutrient content at the stand level.
Internal translocations resulted in changes of concentrations when organs ceased to be physiolog-
ically active and progressively act as sources of nutrients for growth. The stabilized concentrations
observed after translocations took place, were defined as follows:
- needles: initial concentrations were those of the 1-year-old live needles and stabilized concen-

trations were those of the needles in the litterfall.

- woody branches: initial concentrations were those of the youngest branches and final concentra-

tions were those observed on the oldest ones.

- Stem:
A special investigation was made to quantify with sufficient accuracy the internal translocations
in the stemwood as it is the most important component of the stand. Ring width analysis, ring wood
density and ring chemistry were necessary to quantify the amount of nutrients in each ring (in this case
a group of 5 rings) from the sampled trees. It was then assumed that no parameter other than transloca-
tion was involved in the changes of nutrient pool size when rings became older: the translocations were
then estimated from the progressive decrease of a particular ring group nutrient pool size from its initial
development in the younger stand to its final content in the oldest stand.
A subsample of 5 trees among those used for nutrient content evaluation was selected in order to
keep a good representation of girth class distribution: the groups of 5 rings were isolated and measured
in the disks taken each 3 m on the stem of the 40 and 60 year-old stand and each 1.5 m in the 20-year-
old stand. Each individual sample was then analysed for total chemical composition and wood density.
Wood density was determined from fresh volume (water displacement of a saturated sample) and dry

Acta (Ecologica
Nutrient dynamics in a forest stand 77

mass after drying during a week at 100~ (550 samples were analysed for wood density and total
chemistry).
Litterfall was evaluated from 20 litter traps per stand collected every three months.
Biomass and nutrient removal during thinning operations were evaluated from the manager's data
and present relationships established for biomass and nutrient content according to the age of the matter.
The fluxes of nutrients leached from the canopy (crown leaching) used to calculate the nutrient
uptake of the stand were obtained by measuring rainfall and throughfall during the period of observa-
tion (1992-1995) (MARQUES& RANGER, 1996).
Total and "available" soil nutrient reserves were evaluated from soil horizon thickness, soil bulk
density, horizon stoniness and chemical analysis. Samples were collected in 7 individual pits and anal-
ysed separately.

Identification of the main fluxes of the biological cycle

The concepts and methods have already been described by RANGER& COLIN-BELGRAND(1996) in
a study concerning chestnut stands.
- the total requirement of a stand for annual biomass production (woody biomass and foliage) is
the total amount of nutrients in the current annual biomass increment, evaluated when the maximum
annual biomass production is achieved and when the concentration of nutrients has stabilized. Foliar
biomass varied greatly with time, but it is not possible to evaluate these fluctuations with the destruc-
tive sampling method used here. Litterfall gave an index of the variability of the annual needle produc-
tion and was used instead of foliar biomass to evaluate the mean annual requirement of the stands.
- the nutrient uptake of stands from the available soil pool is the sum of nutrients definitively
fixed in the biomass (immobilization) and the nutrients returned to the soil in solid form (litterfall) or in
solution (nutrients leached by rainfall from crown). Uptake of nutrients by a forest stand cannot be
measured directly but can be calculated from a conceptual compartment and flux model described by
RANGER & BONNEAU(1984):
Uptake = Immobilization + Returns (litterfall and crown leaching) _+ (Af + At)
Af is a term used in coniferous stands where foliar biomass is not stabilized. Af must be considered
when the litterfall does not directly correspond to the current needle production because of the delay
between foliar production and litterfall on sempervirent species. This is particularly relevant in very
young stands where crown development is not completed. On the contrary, in old declining stands the
litteffall can be greater than current needle production and Af is then negative. The same applied to
roots, but this compartment was not considered in this study and Ar = 0.
- total internal nutrient translocation from old physiologically-inactive to young growing tissues
is estimated from the difference between the amount of nutrients required for the annual growth of a
stand (requirement) and the amount of nutrients definitively fixed in the woody biomass (immobiliza-
tion) or returning to the soil as litterfall or crown leaching. Translocations that can occur during the
growth period cannot be considered by such a calculation.
- translocation from foliage is the difference between the quantity of nutrients in the foliage
of a stand and the quantity returning to the soil as litterfall or crown leaching.
translocation foliage = (foliage biomass* x concentration of leaving leaves)
- ( f o l i a g e biomass x concentration of litterfall) - crown leaching
(*considered as litterfall biomass in this study)
- translocation from branches: during stand development, the size of the branch compart-
ment increases to reach a specific dimension which depends on the tree species and on silvicultural

VoL 18, n ~ 2 - 1997

78 J. Ranger et al.

treatment. In adult stands, branch increment is compensated for exactly by an equivalent mortality and
the biomass of the woody part of the crown remains constant. The annual increment of branches was
considered for each stand of the chronosequence as the sum of the initial branch quantity (i.e. from the
younger stand) and the increment of branches observed on the immediately older stand, weighed to the
difference of age between these two stands. Translocation from this compartment was therefore, for
each stand, the current increment of the mass of branches multiplied by the difference of concentration
between young branches (which have only active tissues) and old ones (which have a rather stabilized
concentration because translocation still occurs and because young tissues represent a small proportion
of the biomass as the current increment was limited).
translocation branches = Production of branches
• (concentration of young branches - concentration of old branches).
- translocation from stems (stemwood and stembark):
- stemwood: stemwood biomass accumulated strictly with time, and so each nutrient

can be qualitatively and quantitatively studied from its incorporation (when the tissue was active) to its
current situation (when the tissue was no longer active). Translocation was calculated step by step
knowing the ring-group biomass and its nutrient pool size when it was active and when it progressively
started behaving as a source of nutrients withdrawn to young growing organs (fig. 1).
- stembark: translocation from stembark was evaluated assuming that : i) desquama-

tion was negligible and ii) translocation was proportional to that observed for stemwood. This is the
most simple assumption that can be made which of course needs to be checked by more intensive mea-
surements using microanalysis techniques due to the very thin annual increment of bark. Nutrients can
be leached by stemflow from the outer part of stembarks, but it is impossible to quantify this process
and to take it into account in the calculations.
- n u t r i e n t return to soil from litterfatl and crown leaching:
- litteffall was evaluated from traps (see methodology).
- crown leaching was estimated from the throughfall measured on the site since 1992.

Crown leaching was considered to represent 0, 0, 90, 20 and 50%, respectively for N, P, K, Ca and Mg
measured on the throughfall (DAMBRINE& PREVOSTO, 1988).

3 ring group [0-20]

| ring goup [20-40]
]! ring group I40-60]

1 20->60 I

FIG. 1. - Flow chart diagram to estimate nutrient translocation in stemwood.

Acta (Ecologica
Nutrient dynamics in a forest stand 79

- immobilization of nutrients in the biomass is the amount of nutrients definitively fixed when
internal translocation has been achieved. For a particular stage of development of the stand, immobili-
zation is the woody biomass production multiplied by a stabilized concentration i.e. residual concentra-
tion after translocation,

RESULTS

B i o m a s s p r o d u c t i o n of s t a n d s (table III)
T h e p r o d u c t i o n d y n a m i c s of the stands s h o w e d that the m a x i m u m annual incre-
m e n t o c c u r r e d b e t w e e n 20 a n d 4 0 y e a r s b u t t h a t it r e m a i n e d h i g h u n t i l 60.
T h i n n i n g s r e p r e s e n t e d 1 9 % o f t h e s t a n d i n g c r o p b i o m a s s in the 4 0 - y e a r - o l d s t a n d
a n d 14% in t h e 6 0 - y e a r - o l d stand.

TABLEIII.- Dynamics of stand production.

0-20 years 20-40 years 40-60 years

Stem volume production m 3 ha-1 yr-1 8.5 20.8 17.0

total with thinnings no thinnings 24.8 19.4
Stem biomass production t ha-1 yr-1 3.3 7.9 6.4
total with thinnings no thinnings 9.4 7.3
Total woody production t ha-t yr 1 4.1 8.3 7.7
total with thinnings no thinnings 9.9 8.8

D i s t r i b u t i o n o f n u t r i e n t s in the v e g e t a t i o n

A c l a s s i c a l d i s t r i b u t i o n o f n u t r i e n t s w a s o b s e r v e d in t h e s e D o u g l a s - f i r s t a n d s . T h e
C o n c e n t r a t i o n o f e l e m e n t s f o l l o w e d this o r d e r ( t a b l e IV):
young needles > old needles > twigs > b r a n c h w o o d > stembark > s t e m w o o d

TABLEIV. - Mean concentration of major nutrients in the various components of the 60-years-old stand
(data in % of Dry Matter).

N P K Ca Mg
Branches
needles 1 year 1.52 0.12 0.51 0.49 0.12
0.14 0.02 0.12 0.09 0.02
needles > 1 year 1.40 0.09 0.44 0.90 0.12
0.14 0.01 0.10 0.21 0.03
branchwood 1 year 1.14 0.14 0.45 0.49 0.11
0.13 0.02 0.07 0.11 0.03
branchwood > 1 year 0.42 0.05 0.21 0.46 0.04
0.13 0.02 0.08 0,12 0.02
stem
stemwood 0.061 0.003 0.023 0.029 0.005
0.004 0.0001 0.003 0.004 0.0006
stembark 0.33 0.042 0.22 0.309 0.032
0.023 0.006 0.046 0.111 0.006
x.xx = mean value ; x.xx = standard deviation,

Vol. 18, n ~ 2 - 1997

80 J. Ranger et aL

Ca sometimes showed higher concentrations in the old organs than in the young
organs.
The analysis of variance showed that the 40-year-old stand was similar to the
20-year-old stand as far as crown components were concerned and to the 60-year-old
one regarding stem components (RANGERet al., 1995).
The nutrient concentrations used to calculate the translocations from the needles
and branches were selected using criteria defined in the methodology. For the stem-
wood, a more complex investigation was carried out in order to identify the dynamics
of nutrient concentration changes with time. Models were calculated to relate the con-
centration of the ring group to ring parameters like vertical and radial position, local
increment and tree parameters (tree height and tree form factor). Satisfactory common
models for the 3 stands were obtained if explanation of the variance (r2) and residues
(RMSE) were considered, except for P which requires a specific stand model
(Mor~sTn~R, 1993) (table V).
Figure 2 illustrates the mean distribution of N for the 5 trees of the 60-year-old
stand: no major discrepancies occurred between the observed and calculated values.
The general and local variations were restituted by the model. The model was also
quite satisfactory for individual trees.

Annual nutrient requirements of the stands (table VI)

Stand nutrient requirements were mainly linked to tree crown development. Nee-
dles were generally the first nutrient sink and branches generally followed. Stembark
which had a smaller biomass production compared to stemwood, required roughly the
same amount of N and Ms, but had much higher requirements for P and Ca and very
much higher requirements for K. More detailed sampling on stembark is necessary to
obtain as much accuracy as for stemwood.
It was more commonly observed that the demand for stemwood production was
always lower than 10% of total requirements, and even more frequently lower than
5% (60% of the cases).
The effect of age concemed a decrease in the foliar demand with time, an
increase in the demand for branch development with time and an increase in the
demand for stem development parallel to current production. The major difference
concerned the young stand with high requirements linked to needle build-up.

Nutrient uptake to soil (table VII)

Nutrient uptake from soil reserves was strongly linked to the proportion of lig-
neous components and foliar biomass according to stand age. The latter component
always dominated for all nutrients. More generally, the proportion of nutrient uptake
for building up woody biomass increased with time parallel to biomass increment
dynamics.
The total nutrient uptake decreased with stand age for all elements while current
increment strongly increased in the 40-year-old stand and remained high in the oldest
stand. The main causes were the decrease in litter production and a significant
increase in nutrient efficiency to produce biomass with ageing.
Immobilization of nutrients in the woody parts of the vegetation was low when
compared to requirements or uptake. Nevertheless, this is the actual amount of nutri-

Acta (Ecologica
Nutrient dynamics in a forest stand 81

TABEEV, - - Model used to describe the distribution of nutrients in stems.

I. NITROGEN: common model for the 3 stands
N= 130 + 1~1 * HEIGHT+ [~2 * A f +83 * log(fA) + ~4 * CA + ~J5 * CA2 + [~6 * CA 3 + [~7 * AGE + ~8 * AGE2
+ 69 * REHERG + 1310 * REHERG~+ [111 *TREEFF + [~12 * LEVEL2 + e

R2 0.59 adjusted R 2 0.58 Root MSE 8.8

I1- PHOSPHORUS: specific model for each stand

Stand 20-year-old

P = ~ I * f ~1 + 62 * TREEFF+ [J3 * AC3 + 1~4 * LOG(fA ) + 65 * REHETREE + e

Stand 40-year-old
1
P = 61 * f ~ + 62 * LEVEL2 + 63 * REHETREE + ~4 * f A2 + 65 * LOG(fA ) + 66 * fA +

Stand 60-year-old
1 1
P=I30 + 1~1 * ~--~+ 62 * C A 3+ 63 * I N C R E M E N i , + [54 * C130 + 65 * CA 2 + 66 HEIGHT + 67 *

INCREMENT2+

Stand age 20-year-old 40-year-old 60-year-old

R2 0.91 0.801 0,81

adjusted R 2 0.91 0.80 0.80

Root MSE 14.8 11.8 10.5

III- POTASSIUM: common model for the 3 stands

K=60+131 * f ~1+ 1
1~6" ~-'~+ 67 *CA4 + 62 * RESOW + O5*HERIGR2 + ~3*INVRAD + 64*TREEFF

R2 0.89 adjusted R 2 0.89 Root MSE 7.2

IV- CALCIUM: common model for the 3 stands

Ca=130 + 1~1 *fA + 6 2 * f A 2 + 6 3 * f A 3 + ~4,CA 2 + I~5*CA3 + 66*INVRAD + 137*HERIGR +
68*HERIGR 2 + 1~9"RESOW+ 610*REHERG

R2 0.76 adjusted R 2 0.76 Root MSE 6.4

V- MAGNESIUM: common model for the 3 stands

Mg = 130 + [~1 * log(fA) + 62 INVRAD + 133 * HAUTEC2
+ ~4 * RESOW + 65 * f A 2 + 66 * TREEF + ~7 * TREEF2 + ~8 * AGE + 69 * C1302 + ~10 *RAD2 + ~11
1
9INCREMENT + e

R2 0.75 adjusted R 2 0.74 Root MSE 1.4

CA: cambial age of the ring (counted from the pith) (years); f A : physiological age of the ring (counted from
the bark) (years); age: tree age; height: total tree height (m); herigr: height of the ring group from the soil
(absolute value, m); increment: ring group width (mm); level: level of the sampled disk (m); tad: tree radius;
reherg: relative height of the ring group; rehetree: relative position of the sample in the stem; resow: softwood
proportion; treeff: tree form factor.

Vol. 18, n ~ 2 - 1997

82 J. Ranger et al.

TABLEVI. - Current annual requirements of nutrients for biomass increment

in the three stands of Vauxrenard (Beaujolais).

Dry
age comparmaent Matter N(1) N% P(1) P% K(1) K% Ca(l) Ca% Mg(l) Mg%
(1)
20-year- Needles 4100 65.6 82 4.5 53 23.8 35 19.3 77 4.9 76
old stand Branches 800 8.8 11 1.0 12 4.5 7 3.4 13 0.9 14
Stemwood 2845 1.7 2 0.2 2 1.6 2 1,1 4 0.2 3
Stembark 430 3.7 5 2.7 32 38.2 56 1.4 5 0.5 7
TOTAL 8175 79.8 100 8.4 100 68.1 100 25.1 100 6.5 100
40-year- Needles 2600 42.6 68 2.9 42 13.8 42 10.9 51 3.1 56
old stand Branches 1140 12.8 20 1.5 21 5.7 17 4.5 21 1.4 25
Stemwood 6895 3.7 6 0.4 6 3.3 10 2.2 10 0.4 8
Stembark 1005 3.7 6 2.2 31 9.9 30 3.9 18 0.6 11
TOTAL 11640 62,8 100 6.9 100 32.7 100 21.4 100 5.5 100
60-yem- Needles 2500 38,0 53 3.0 33 12.8 30 12.3 35 3.0 45
oldstand Branches 2375 27.l 38 3.3 36 10.7 25 11.6 33 2.6 39
Stemwood 5610 3.4 5 0.4 4 2.8 6 1.9 5 0,3 4
Stembark 815 3.2 4 2,5 27 16.7 39 9.4 27 0.8 12
TOTAL 11300 71.7 100 9.2 100 42.9 100 35.2 100 6.7 100
(1) = data in kg per ha.

ents which disappeared from soil reserves. Tree crown participation (woody branches)
decreased with stand age but remained as high as 15% for N, 35% for P, 46% for K,
30% for Ca and 70% for Mg in the 60-year-old stand.

TABLEVII, - Evaluation of the mean annual uptake as a function of stand age

(data in t per ha for biomass and kg per ha for nutrients).

Stand age Dry matter N P K Ca Mg

20-year-old stand Immobilization 4.1 7.4 0.7 4.5 11.0 0.7
Litterfall 4.1 59.6 4.3 6.8 37.8 4.1
Leaching 0.0 0,0 25.1 0.8 0.8
Uptake 67.0 5.0 36.4 49.6 5.6
40-year-old stand Immobilization 8.3 5.9 0.5 2.8 - 3.2 0.5
Litterfall 2.6 25.1 2.8 3.7 14.9 2.1
Leaching 0.0 0.0 11,0 1.4 0.7
Uptake 31.0 3.3 17.4 13.2 3.3
60-year-old staud Immobilization 7.7 6.5 0.5 0.5 9.0 0.5
Litterfall 2.5 25,9 1.8 3.6 :t8,8 1,8
Leaching 0.0 0.0 9.5 1:4 0.8
Uptake 32A 2.3 13.6 29.2 3.0
Immobilization:calculated from stabilized concentrations of ligneous compartments for the periods 0-20, 20-40
and 40-60 years.
Litter: measured for the years 1992-1995.
Uptake = Immobilization + Restitution (litterfall + crown leaching).
Crown leaching was considered as 0.0. 90. 20 and 50 % of the net throughfall for N, P. K, Ca and Mg respectively.

Acta ,~.cologica
Nutrient dynamics in a forest stand 83

a - Observed values

' 800

700

6O0

z 2
i 4o 6o 3oo

38 Camblal age (years)

24
30 o
Vertical level (111)

b - Predicted values

6~
Q.

z 2

C~mb:al age (yearsl

30 o
V~'lioal level (m)

Fic. 2. - Mean distribution of N for the five trees of the 60-year-01d Douglas-fir stand:
a) measured values and b) values calculated by the model.

Internal translocations in trees (table VIII)

T h e b e h a v i o u r Of nutrients was v e r y different: N, P, K and M g s h o w e d p o s i t i v e

t r a n s l o c a t i o n w h i l e C a s h o w e d a total n e g a t i v e t r a n s l o c a t i o n w i t h i n the trees, c o r r e -

VOI. 18, n~ 2 - ]997

84 J. R a n g e r et al.

sponding to a net accumulation in older tissues. N translocation was higher than that
of the other nutrients.

TABLEV I I I . - Quantification of internal translocation in trees according to hypothesis 11

(data in kg per ha and year).

compartment N N% P P% K K% Ca Ca% Mg Mg%

20-year-old needles 5.9 51.5 0.2 12.6 - 7.9 - 243.8 - 20.4 104.1 0.1 13.9
stand branches 5.4 46.8 0.6 40.4 2.2 69.1 0.6 - 3.3 0.6 77.8
stemwood 0.1 0.6 0.0 2.8 0.4 11.4 0.1 - 0.4 0.0 2.8
stembark 0.1 1.2 0.7 44.2 8.5 263.3 0.1 -0.5 0.0 5.6
TOTAL 11.5 1.6 3.2 -- 19.6 0.7
40-year-old needles 17.2 65.5 - 0.2 - 14.2 - 0.5 - 8.4 - 6.6 149.0 0.3 19.2
stand branches 8.1 30.8 1.0 73.0 3.1 51.8 0.8 - 18.t 0.9 58.1
stemwood 0.3 1.1 0.1 4.3 0.5 8.7 0.3 -7.2 0.1 4.8
stembark 0.7 2.6 0.5 36.9 2.9 47.9 1.1 - 23.7 0.3 17.9
TOTAL 26.3 1.4 6.0 - 4.4 1.6
60-year-old needles 10.6 40.4 1.2 15.8 - 0 . 3 5.5 -9.5 414.8 0.4 2000.0
stand branches 17.1 65.2 2.1 28.2 5.7 - 104.3 0.7 - 3 1 . 0 1.7 8300.0
sternwood -0.3 - 1.3 0.1 1.1 1.1 - 19.5 0.4 - 17.0 0.1 600.0
stembark - 1.2 -4.4 4.2 54.9 - 11.9 218.3 6.1 - 2 6 6 . 8 - 2.2 - 10800.0
TOTAL 26.2 7.6 - 5.5 - 2.3 0.0

The various components presented specific behaviour with translocations gener-

ally related to requirements: translocations within tree crown were generally highest,
but stembark also gave high values. The most constant observation was the low abso-
lute values of internal translocations from stemwood when compared to the other tree
components. If cumulated internal translocations in stemwood during the rotation
were compared to the immobilization in the same component (data not shown), the
conclusions would be somewhat different: N was not translocated in stemwood but
translocations of P, K, Ca and Mg represented 44, 60, 17 and 30% respectively of
their immobilizations ( B E R N A R D , 1994).
Translocation efficiency tended to increase with stand age for all elements
excluded K and Mg in the 60-year-old stand.

DISCUSSION

Dynamics of the main fluxes of the biological cycle

Stand requirements did not vary much with stand age even if they tended to be
higher in the young stand, especially when compared to the 40-year-old stand
(table IX). This is linked particularly to crown development as stand immobilization
did not vary greatly with time. Litterfall indicated the same trend, as it was used
instead of needle biomass in the hypotheses retained for calculation.
The majority of the nutrients taken up from the available soil pool for plant nutri-
tion returned to the soil forest floor with litterfall. K showed a specific behaviour with
large crown leaching flux representing more than half of the young stand K require-
ment. Direct uptake of ammonium by tree crown from atmospheric deposits explained

Acta (Ecologica
Nutrient dynamics in a forest stand 85

TAaLE IX. - Evaluation of the current annual fluxes according to stand age (data in kg per ha and year).

age flux N P K Ca Mg
20-year-old Requirement 79.8' 8.4 68.1 25.1 6.5
stand Leaching 0.0 0.0 25.0 0.8 0.8
Immobilization 7.4 0.7 4.5 4.6 0.7
Litterfall 59.6 4.3 6.8 37.8 4.1
Uptake 67.0 5.0 36.4 43.2 5.6
Translocation 11.5 1.6 3.2 - 19.6 0.7
40-year-old Requirement 62.8 6.9 32.7 21.4 5.5
stand Leaching 0.0 0.0 11.0 1.4 0.7
Immobilization 5.9 0.5 2.8 3.3 0.5
Litterfall 25.1 2.8 3.7 14.9 2.1
Uptake 31.0 3.3 17.4 19.7 3.3
Translocation 26.3 1.4 6.0 - 4.4 1.6
60-year-old Requirement 71.7 9.2 42.9 35.2 6.7
stand Leaching 0.0 0.0 9.5 1.4 0.8
Immobilization 6.5 0.5 0.5 10.1 0.5
Litterfall 25.9 1.8 3.6 18.8 1.8
Uptake 32.4 2.3 9.5 29.2 3.0
Translocation 26.2 7.6 - 5.5 - 2.3 0.0

this heavy K leaching in the young stand by a simple ion-exchange reaction (MARQUES
& RANGER, 1996).
The proportion of nutrients required for stand development, as taken up from soil
reserves, decreased with time. On the contrary, the proportion of nutrients translocated
from old tree components to young developing ones tend to increase with time espe-
cially for N and P. Ca behaved rather differently with negative translocations indi-
cating that it continued to accumulate during senescence (BRINGMARK, 1977;
HELMISAARI,1992; COLIN-BELGRANDet al., 1996). This clearly indicates that, apart for
Ca, the forest stand acquired a relative independence from soil nutrient reserves with
time: internal translocations of major nutrients within the trees represented a key pro-
cess in the ecosystem, increasing the efficiency of the limited pool of soil nutrients to
produce biomass.
From these results (table X) a simple scheme can be proposed for the mineral
functioning of N, P, K and Mg for this forest stand:
- in the young stand: for 100% requirement, uptake from soil ranged from 55 to
85% depending on elements (except Ca), and internal translocations ranged from 5 to
20%; immobilization was no more than 20% of the requirements as more than 70%
of uptake returned to the soil through litterfall or crown leaching (for "soluble" ele-
ments like K).
- in the older stand: for 100% requirements, uptake from soil ranged from 20 to
45% (except Ca). The internal translocation remained high for N and P and the immo-
bilization was still very low (<10% of requirements). Litterfall always represented a
large proportion of the elements taken up from the soil and then strongly contributed
to maintaining a large amount of nutrients in the soil top layer.
These results are in agreement with those of SwrrZER & NELSON (1972),
M~KONEN (1974), BRINGMARK(1977), BORMANN& LIKENS(1979), MILLER(1981),

Vol. 18, n ~ 2 - 1997

86 J. Rangeret al.

TABLE X. - - Cumulative fluxes in the three stands: relative values.

N P K Ca /rig
20-year-old stand Uptake as a % of requirements 84.0 59.9 53.5 172,1 86.2
Translocation as a % of requirements 14.4 19.0 4.7 - 78.1 10.8
Litterfall as a % of requirements 74.7 51.2 10.0 150.6 63.1
Immobilization as a % of requirements 9,3 8.7 6.6 18.3 10.8
Translocation as a % of uptake 17.2 31,8 8.8 - 45.4 12.5
Litterfall as a % of uptake 89,0 85.5 18.7 87.5 73.2
40-year-old stand Uptake as a % of requirements 49,4 48.4 53.2 92.1 60.0
Translocation as a % of requirements 41.9 20.3 18.3 - 20.6 29.1
Litterfall as a % of requirements 40,0 40.6 11.3 69.6 38.2
Immobilization as a % of requirements 9,4 7.8 8.6 15.4 9.5
Translocation as a % of uptake 84.8 41,9 34.5 - 22.3 48.5
Litterfall as a % of uptake 81.0 83.8 21.3 75.6 63.6
60-year-old stand Uptake as a % of requirements 45.2 25.0 22.1 83.0 44.8
Translocation as a % of requirements 36.5 82.6 - 12.8 - 6.5 0.3
Litterfalt as a % of requirements 36.1 19.6 8.4 53.4 26.9
Immobilization as a % of requirements 9.1 5.4 1.2 28,7 7.5
Translocation as a % o f uptake 80,9 330.4 - 57.9 - 7.9 0.7
Litterfall as a % of uptake 79.9 78.3 37.9 64.4 60.0

KHANNA• ULRICH(1991) or HELMISAARI(1990) and enable a real understanding of

mineral cycle at the ecosystem level.

Nutrients involved in forest growth during rotation

Cumulation of fluxes during rotation is a rather theoretical way to present the
data but is very useful i) to showing the real amounts of nutrients involved in tree
nutrition, ii) to describe the efficiency of the system in using the limited pool of nutri-
ents to produce biomass and iii) to point out the relevance of the biological cycle in
the soil function,
The results indicated that about 4300 kg of N, 500 kg of P, 2900 kg of K, 1600 kg
of Ca and 400 kg of Mg were required for biomass production, which was enormous
when compared to current available resources presented in table II.
When looking in more detail at the 60-year-old stand which represents more or
less the mean rotation length for Douglas-fir in this area, it appears that 2610 kg of
N, 210 kg of P, 1270 kg of K, 1850 kg of Ca and 240 kg of Mg were extracted from
soil reserves during rotation. This represents 59%, 236%, 309% and 190% of the
available P, K, Ca, Mg reserves respectively and 35% of the total N reserve.
This key parameter must be determined because nutrient uptake from soil
reserves represents the amount of elements which are absorbed by plant roots on all
the rooting zone. It will be of paramount importance in the future to have a better
understanding of the relationship between nutrient uptake and soil nutrient avail-
ability~ It has been postulated that the current stock of available nutrients must be
50 times greater than the uptake of nutrients strongly fixed in the soil like P, and only
10 to 20 times for nutrients less strongly fixed like Ca and Mg for a non limiting
uptake; K seems to have an intermediate position (RAN~ER & BOWNEAU, 1984). This
means that the available soil reservoir must contain 250 kg per ha of P, between 400
and 800 kg of Ca, between 60 and 120 kg of Mg and between 500 and 1000 kg of K

Acta (Ecologica
Nutrient dynamics in a forest stand 87

to sustain the annual increment of the stand during all the phases of its development.
Comparison with table II showed that available K, Ca and Mg reserves could become
rapidly limiting if no care is taken in the management of this ecosystem.
For 60 years, atmospheric deposits have provided a significant amount of these
elements namely the equivalent of 42% for N, 45% for P, 15% for K, 18% for Ca and
24% for Mg taken up by the vegetation. On the other hand, large amounts of nutrients
were lost in seepage waters, always larger than atmospheric input (MARQUES &
RANGER, 1996). Only the deposits occurring during the growing season benefited
forest growth because the major part of the winter atmospheric input was drained out
of the ecosystem.
In the 60-year-old stand, the current annual uptake of P, K, Ca and Mg repre-
sented 0.6, 0.9, 3.6 and 1.1%, respectively of the available reserves.
Litterfall and crown leaching brought back to the soil surface more than 80% of
the nutrients taken up by roots in all the soil layers.
The final immobilization is small when compared to the large amounts of nutri-
ents required for forest stand growth. Immobilization represents the real amount of
nutrients substracted from the available soil pool during rotation. The amount of nutri-
ents definitively leaving the ecosystem will vary according to the harvesting intensity
i.e. from debarked stems to whole tree harvesting. Whole tree harvesting will result
in a large depletion of soil nutrients as does the withdrawing or burning of the rem-
nants (LICKENSet al., 1977; ADAMS& BOYLE, 1980; MORRISet aL, 1983). Small immo-
bilization of nutrients in forest crops has surely led to the legendary concept of forest
tree frugality which concealed the real requirements of forests as demonstrated here
in the case of Douglas-fir stands.
These results clearly demonstrate that soil fertility is a dynamic not a static
parameter: the ecosystem will stay in its present equilibrium as long as the input-
output budget of the system will be itself balanced. The flux of nutrients released by
weathering processes fixes the maximum rate of depletion associated with harvesting
operations.
Restitutions by litterfall brought back approximatively 210 t of organic matter
(OM) or 100 t of carbon to the soil upper layer during rotation. They represent about
5 times the forest floor mass at the end of the rotation. This is one of the most impor-
tant impacts of vegetation on soil function because OM plays an important role in soil
fertility: i) it is an important reservoir of nutrients released by mineralization pro-
cesses (see potential reserves in table II), ii) OM is a physico-chemical support for
cation fixation and iii) OM participates in soil aggregation thus conditioning soil
drainage and aeration.
Thinnings representing 15% of the standing crop have to be taken into account
when budgets are established for forest plantations (table XI).
No observation of underground biomass was made in this study. This is not really
very important for large roots, as estimations can be made from literature (SANTAN-
TONIO, 1979; FAHEYet al., 1988; RANGERet al., 1992) even if the dynamics of large-
root biomass production and nutrient incorporation along the rotation is not very well
documented. Biomass of the root-system would represent between 10 and 20% of the
above ground biomass, depending on species treatments and sites. Immobilization

Vol. 18, n~2 - 1997

88 J. Ranger et aL

T~a~LEXI. - Cumulative fluxes for the 60-year rotation of Douglas-fir of Vauxrenard (Beaujolais)
(data in kg per ha and year).

flux MS N P K Ca Mg
Requirement FS 718300 4286 490 2874 1634 374
thinnings % 15 23 23 20 25 24
thinnmgs Th 106593 967 113 578 402 89
FS + Th. FS +Th. 824893 5253 603 3452 2036 463
Leaching FS 0 0 910 72 46
thinnings % 23 23 20 25 24
thinnmgs Th 0 0 183 18 11
F S + Th. FS +Th. 0 0 1093 90 57
Immobilization FS 402000 396 35 156 360 34
thinnings % 14 15 16 20 15 19
thinnmgs Th 55960 61 6 3I 53 7
FS +Th. FS + Th. 457960 457 41 187 413 41
Litterfall FS 184000 2212 178 282 1430 160
thinnings % 15 23 23 20 25 24
thinnmgs Th 27305 499 41 57 352 38
FS + Th. FS +Th. 211305 2711 219 339 1782 198
Uptake FS 2608 213 1266 1842 238
thinnings % 15 16 20 15 19
thinnings Th 403 35 256 270 46
FS + Th. FS +Th. 3011 248 1522 2112 284
Translocation FS 1280 212 74 -526 46
thinnings % 23 23 20 25 24
thinnings Th 289 49 15 - 130 11
F S + Th. FS +Th. 1569 261 89 - 656 57
FS = Final Stand; Th = Thinnings.

would then represent between 15 and 25% of the nutrient content of the aerial woody
biomass.
The main difficulty concerns the dynamics of fine root production which is the
most relevant point for understanding the dynamics of nutrients in the ecosystem. The
second point is very much more difficult to estimate than large-root behaviour.
According to SAyrAbrro~o et al. (1977) for Douglas-fir plantations, between 5 and
10 t of fine-root dry matter were produced each year on mature stands. Large differ-
ences occurred according to site water regimes, but various methodologies can lead
to very different estimations.
It did not seem relevant to take the dynamics of the belowground part of the veg-
etation into account, without any measurements. Nevertheless, one must consider that
at least the same amount of carbon and nutrients as those measured from aerial bio-
mass was cycled from fine-root and symbiont turnover. One major difference with
aerial biomass is that root turnover contributes to local recycling contrary to crowns
which bring back nutrients taken up in all the rooting zone to the soil surface.

CONCLUSIONS

Present soil nutrient reserves are relatively small when compared to stand
nutrient uptake. This means that the biological cycle represents the key process in the

Acta (Ecologica
Nutrient dynamics in a forest stand 89

m a i n t e n a n c e of forest productivity. More precisely, it appears that organic matter min-

eralization and mineral weathering are the parameters determining the level of avail-
able nutrients and consequently forest production.
Forests have to be m a n a g e d not only according to stand i m m o b i l i z a t i o n but also
keeping in m i n d that the a m o u n t of available nutrients will be in agreement with the
bulk uptake of nutrients. Another major point for forest m a n a g e m e n t is to take into
account the role of soil organic matter. The majority of the nutrients taken up from
the soil are brought back to the soil surface as organic c o m p o u n d s which slowly min-
eralize giving a flux of nutrients available for new uptake. Destruction of the organic
layer especially during the regeneration phase represents a major disturbance in eco-
systems with limited nutrient reserves, surely responsible for soil acidification and
decrease in fertility.
New forest m a n a g e m e n t practices will have to take into account the classical laws
of agronomy better than they do at present if sustainable m a n a g e m e n t is to be
achieved (MILLER, 1981).
M o n i t o r i n g a site for a long time or at least having mid term series of measure-
ments on chronosequences of stands will provide the most relevant information for a
better understanding of ecosystem dynamics. This information will make it possible
to validate both ecosystem function models as well as m a n a g e m e n t models and man-
agement tools (DvcK e t al., 1994).

ACKNOWLEDGEMENTS

We would like to thank:

- the research teamof the "Cyclesbiogrochimiques"for theirtechnicalassistance,DominiqueGELHAYEfor
managingthe fieldwork,SrverineBrENAIME,LouisetteGEtaJArEand Beno~tPOLLmRfor laboratorymeasurements.
- the forest managers,the "Office Nationaldes For~ts", Grrard POLYand BernardJOEARDfor providingall
the facilitiesduringinvestigationand sampling,
- Claire GAYand ChristineYOUNG,INRA translationdepartment, for revisingthe English version of the
manuscript,
- the "Minist~rede l'Agriculture,Divisionde l'EspaceRural et de la For&", and the EuropeanUnion (AIR
Research Contract) for financialsupport.

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Acta tTEcologica