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Int. J. Fund. Appl. Sci. Vol. 4, No. 4 (2015) 91-98

Open Access

ORIGINAL ARTICLE
ISSN 2278-1404

International Journal of Fundamental & Applied Sciences

Litter decomposition and nutrient cycling in temperate forest of Kumaun
region
Lovy Raj Chandraa, b *, Apurva Raia, Veena Pandeb and Nandita Singha
Plant Ecology and Environmental Science Division, CSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow226 001, Uttar Pradesh, India; bDepartment of Biotechnology, Bhimtal Campus, Kumaun University, Nainital-263136,
Uttarakhand, India
Manuscript received 24th Sept, 2015, revised 16th Oct, 2015, accepted 26th Oct, 2015

Abstract
Background & Aim: The magnitude of litter fall and its decomposition in a forest ecosystem impart a crucial role in
nutrient supply to plant as well as microorganisms. The present study was conducted to evaluate and compare the
litter fall production, leaves litter decomposition and nutrient re-translocation among the dominant species i.e. Pine
and Oak, of temperate forest of Kumaun region. Results: The estimated annual litter fall production was significantly
23% higher in Pine stand than Oak stand. The maximum portion of litter fall was occurred in summer and dry period
between April-June. The nutrients concentration was higher in green leaves than leaves litter. Additionally, the
nutrients concentration was found to be high in green leaves as well as in leaves litter of Oak species. The
decomposition rate of leaves litter was found to high in Oak floor which is inversely proportional to the C/N ratio.
The nutrient immobilization for all the nutrients was found to be high in Pine stand. The nutrient re-translocation
percentage were in the order of P > K > N > Ca, in both the species that indicates the higher immobilization of
phosphorous and potassium in the region. Soil status is also responsible for litter decomposition. The soil MBC,
MBN, MBP, basal respiration and enzymes activity were higher in Oak stand which is directly correlated with the
decomposition rate. Conclusions: Our results suggested that different plant species growing in same climatic
condition differed in litter fall, litter decomposition and have different level of nutrients uptake and availability.
Keywords: Litter decomposition; Nutrient cycle; retranslocation; Pinus roxburgii; Quercus leucotricophora
@2015 BioMedAsia All right reserved

1. Introduction
The energy required by various ecosystem processes is
mainly controlled and available from photosynthesis.
Soil does not have capability to capture energy directly
from sun i.e. solar energy because of the lack of
photosynthetic organisms and depends on the energy rich
bodies mainly produced by plants residues. These plant
residues are decomposed by a series of naturally
occurring biological processes that involve both
microbial flora and fauna. Litter fall from leaves and
other part of plants is the basic character of all vegetative
environments, the vital source of nutrients and the
beginning point of nutrient cycle. It is the main pathway
through which organic matter and nutrients returns to the
soil surface, where it composes the major portion of
nutrients available to soil, incorporation with roots
*Corresponding author
Full Address :
CSIR-National Botanical Research Institute, Rana Pratap Marg,
Lucknow-226001, India
Phone no. +91522-2205931
E-mail: [email protected]

turnover1, 2. The availability of nutrient to the soil reflects

the rate of decomposition and microbial activity3. Litter
fall along with roots composes a major segment of
nutrient cycling between plants and soils, thus internal
fluctuations of C, N and P at ecosystem level are
constrained by the available litter fall4, 5. The nutrients
availability in any given soil system is mainly because of
the decomposition dynamics of organic matter. The
major part of carbon and the energy that enters the food
web is detritus based litter that are used by the
heterotrophic organisms and act as a nutrient reservoir in
the food web. Plant litter production and decomposition
is a crucial ecosystem process that defines and governs
the plant-soil relationships by regulating the nutrient
turnover and the buildup of soil organic matter. In fact,
recycling of nutrients through decomposition is the
primary source of available nutrients for plants in most
unmanaged terrestrial ecosystems6. In contrast, the
ecosystem existence is based on the decomposed matter.
The rate of decomposition and the nutrient releases are
highly influenced by the tree species through different
properties, such as magnitude of litter produced, litter
91

Chandra et al.

quality and nutrients release as well as climatic

conditions and existing microbial communities in the soil
system7.
Forests worldwide are known to be significantly
important habitats acts as a storehouse of biodiversity.
These forests are the source of a major part of global
terrestrial carbon in which temperate forest ecosystem
plays major role in carbon sequestration from increasing
atmospheric carbon dioxide, as it covers the major
portion of terrestrial land. The productivity and the
functioning of a forest are mainly influenced by the
pattern and the rate of nutrient cycling. In a forest
ecosystem, litter fall and its decomposition act as a
clogged system which evidently interrelated with the
cycling of nutrients; it regulates the nutrients and energy
flow between biological components and soil, fixes
energy and governs the overall growth and productivity
of forest8, 9.
The magnitude of nutrient re-translocation and the
nutrient use efficiency in litterfall is the key source of
nutrients for plants as well as used as an indicator of soil
nutrient status10. Along the same climatic conditions,
different plant species respond differently in their
nutrient release pattern. The re-translocation of nutrients
and their movement via soil and leaf litter is
characterized as one of the important process in nutrient
dynamics that are prominently used by plants to conserve
nutrients.
Numerous studies have been documented on various
features of litter fall in different forest ecosystem all over
the world as well as in India. However, the information
regarding different aspects of litter fall for temperate
forest of India is still limited. The present study is
undertaken in Nainital district of Kumaun region located
in Uttarakhand. The forest covers a diverse range of
plantation11. The main objective of the study is to
examine the ability of two dominant plant species of the
region in their litter fall creation and litter decomposition
under same environmental condition and their impact on
nutrients availability and uptake.

Int. J. Fund. Appl. Sci. Vol. 4, No. 4 (2015) 91-98

of the region has been described by Singh and Singh,

1986 12. The climate is summer monsoon with three
distinct season categorized into summer, which is warm
and dry (March-May), rainy season is warm and wet
(June-September) and the winter is cold and moderately
dry (October-February). The minimum temperature
ranges from -3 C to 12 C during the month of January
and the maximum temperature fluctuates from 18 C to
30 C in the month of June. The average annual rainfall
is 151.9mm (Figure. I). The soil is generally brown in
color with 58.22%, 11.33% and 30.45% of sand, slit and
clay respectively.
2.2 Litter fall collection
To study litter input in the selected forest stands i.e. Pine
and Oak forest, five litter traps were randomly placed in
each stands within the established plots of 10X10m2. The
litter accumulated within the traps was collected at
regular interval time period for two consecutive years i.e.
Oct, 2012 to Sep, 2014. Litter samples were taken to the
laboratory oven dried at 60 C and then weighed. The
green leaves/needle samples of Pine and Oak were also
collected at the same time.
2.3 Chemical analysis
The dried leaf litter and the green leaves samples were
then processed for chemical analysis. The carbon content
was measured by loss of ignition method. The nitrogen
content was determined using Kjeldahl method13. Carbon
content was measured by loss of ignition method.
Stannous chloride method14 followed by hot plate
digestion in HNO3:HClO3 (3:1) at 180 C for 6 h was
used for phosphorous and potassium concentration.
2.4 Litter decomposition
The litter decomposition rate was estimated using nylon
bag technique. A total of 50 litter bags (25 bags for each
stand) were prepared. Each Litter bag of 20X20cm (5mm
mesh) was filled with 10gm of air dried weighed of litter
sample and placed on the forest floor in October, 2012.
The litter bags were drawn at fixed interval of time
during the study period. After collection, the extra

2. Methods
2.1 Site description
The study was conducted in the unprotected forest of
Nainital district, located in Uttarakhand. The pine (Pinus
roxburgii) and oak (Quercus leucotrichophora) are the
two most dominated plant species of the region. The
selected area is divided into two stands on the basis of
dominant plant species. The stand dominated with pine
plantation is surrounded between E 079 32' 22.9", N 29
23' 24.4" and at elevation of 1822m. Similarly, oak
dominated stand is lie within E 079 33' 08.3", N 29 21'
15.9" and at elevation of 1333m. The vegetative structure
92

Figure I: Rainfall and temperature from Oct, 2012 to Sep,

2014 in the experimental site (Source: India Meteriological
Department Site)

Litter decomposition and nutrient cycling in forest of Kumaun region

residual material was removed and the wet weight of the

remaining litter sample was measured. The samples were
then oven dried at 60 C and reweight until constant
weight achieved.

Int. J. Fund. Appl. Sci. Vol. 4, No. 4 (2015) 91-98

k = Decomposition rate constant

t = Elapsed time (year-1)
The nutrient re-translocation efficiency (NRE) was
computed by the equation given by Finzi et al.27.

2.5 Soil study

NRE % = {(A B) / A} 100

Soil samples were randomly collected from each plot at a

depth of 0-10cm and were taken to laboratory for
analysis. The samples were then air dried, grounded and
sieved using 2mm mesh screen. Soil pH was measured
by the Orion star ion analyzer using soil:distilled water
(1:2.5). Total organic carbon was analyzed using method
proposed by Walkey and Black method15. Available and
total nitrogen was estimated using Stanford and smith16
and Kjeldahl method, respectively. Available and total
phosphorous was assessed using Olsen method17 and
stannous chloride method14. Available potassium was
determined by flame photometric method18.
Soil microbial biomass carbon, nitrogen and
phosphorous (MBC, MBN and MBP) was estimated
using chloroform fumigation extraction method. Final
MBC, MBN and MBP were calculated by difference
between fumigated and non-fumigated values with a
conversion factor of 0.33 for MBC, 0.54 for MBN and
0.40 for MBP.
Soil basal respiration was determined by alkali
absorption method that quantifies CO2 evolution using
moist soil sample19. Soil enzyme assay were perform in
moist soil. Acid phosphates and -glucosidase activity
was measured using Eivazi and Tabatabai 20, 21.
Dehydrogenase activity was assayed by Pepper et al. 22
method. Protease activity was estimated using Ladd and
Butler 23 protocol. Fluorescein diacetate (FDA) activity
was measured using method proposed by stubberfield
and Shaw24. Cellulase activity was quantified using
Schinner and Von Mersi method25.
2.6 Data analysis
The remaining dry mass for each period was calculated
using the weight of litter at each sampling period and the
initial weight using the formula:
% RM= (Wt/W0) X 100
Where, Wt = Weight (Wt) of litter at each
sampling period (t)
W0= Initial mass
Decomposition rate constant (k) of each single species
was calculated by the most widely used single
exponential decay model from the changes of litter dry
mass over time26:
ln (Wt/W0) = -kt
Where, Wt = Amount of litter at time t
W0 = Initial mass of litter

Where, A= Nutrient in green leaves

B= Nutrient in leaf litter
Nutrient use efficiency (NUE) was calculated
according to Vitousek28.
NUE=Litterfall mass/Nutrient content in litterfall
The values were expressed in g m-2 yr-1.
Data were summarized as mean SD (standard
deviation). Values were analyzed by Students t test to
determine the significant difference among the two
stands in term of litterfall production, decomposition rate
and the nutrient uptake. Soil samples were also analyzed
for the significant difference among the two stands. A
two-tailed (=2) p value less than 0.05 (p < 0.05) was
considered to be statistically significant.

3. Results and discussion

3.1 Litter fall
Under the same climatic condition, the composition of
tree species plays an important role in litter production29.
The two plant species found to be different in the
quantity of litter biomass production. The mean annual
litter fall biomass was 687.22 and 527.74 g m-2 y-1 in
Pine and Oak, respectively (Figure. IIb). A significant
difference (p<0.05) was found among the litter fall
production by the two species. The litter fall at Pine
stand is computed 23% higher than Oak stand. The
maximum portion of litter fall biomass was observed in
summer during April-June (Fig. IIa), which is
characterized as dry period, and the result is in agreement
with others result30, 1. To have a better understanding of
nutrient cycling, forest growth, successional pathways
and interactions with environmental variables in forest
ecosystems the evaluation of litter fall production is
needed. Climate, season, substrate quality and type of
vegetation vary with varying litter production31, 32.
3.2 Nutrient Status
The nutrient index in green leaves and leaf litter were
found to be significantly different (p<0.05) in both the
species. The nutrient concentration of N, P, K and Ca in
green leaves and leaf litter were outlined in (Figure. IIIa).
The variation in nutrient quality is mainly because of
varied plant species differ in nutrient quality. The
concentration of P, K and Ca in green leaves of Pine was
recorded 0.54 mg g-1, 2.15 mg g-1 and 12.14 mg g-1,
wheras the values in leaf litter were 0.32 mg g-1, 1.37 mg
g-1 and 8.69 mg g-1, respectively. Similarly, the P, K and
93

Chandra et al.

Int. J. Fund. Appl. Sci. Vol. 4, No. 4 (2015) 91-98

Figure II: (a) Litterfall pattern and (b) Total Litterfall in Pine and Oak stands (g m-2 yr -1)

Figure III: (a) Nutrient concentration in green leaf and leaf litter of two plant species (mg g-1)
(b) Percentage of remaining mass at Pine and Oak stands during the study period

Ca concentration in Oak was 0.99 mg g-1, 3.56 mg g-1

and 14.20 mg g-1 for green leaves and 0.58 mg g-1, 2.29
mg g-1 and 10.36 mg g-1 for leaf litter, respectively. The
nitrogen (%) in Pine was recorded 0.025% and 0.018%
in green leaves and leaf litter. For Oak, the % nitrogen
was found to be 0.030 and 0.024 in green leaves and leaf
litter. Additionally, the Oak litter found to be low C/N
ratio than that of Pine litter i.e. 16.78 and 21.84. This C/
N ratio is a good indicator of decomposition process. A
significant variation (p < 0.05) has been observed among
all the mobile nutrients between leaf litter and green
leaves. The concentration of all the nutrients was found
to be low in leaf litter than the green leaves in both the
plant species. The two plant species have different traits
displayed species-specific variation in nutrient pattern.
The plant species induced variation in chemical
composition which directly affects the microbial
attributes residing in that particular soil and influences
the decomposition of organic matter as well as the
physico-chemical status of soil3, 33.
3.3 Litter decomposition
The rate of decomposition is highly influence by litter
quality. The litter decomposition was found to be
positively correlated with the time elapsed (r = 0.98, p <
0.05). The mass remained at the end was 47.6% and
36.2% for Pine and Oak, respectively (Figure. IIIb). The
calculated annual decomposition rate of Pine and Oak
94

litter was 0.74 and 1.01. The Pine litter has

comparatively slower rate of decomposition than the Oak
litter residing under same climatic condition. Pine litter
with low nitrogen content and high C/N ratio results in
significantly slower decomposition than Oak litter, that
ultimately responsible for low weight loss % at Pine
stand. The results are adjacent with other studies which
have reported significantly positive correlation between
C/N ratio and litter decomposition34, 35. The difference in
the litter quality and the decomposition rate between the
two stands dominated with two different plant species is
mainly due to the difference among the substrate quality.
Few literatures have demonstrated that stands with
higher nutrient pool exhibit faster decomposition of
litter3. Additionally, the efficient nutrient use efficiency
is primarily described by lower nutrient concentration in
litterfall28. The nutrient use efficiency of the selected
species specific-stands of temperate forest of Kumaun
region is presented in (Fig. IVa). The nutrient i.e. N, P,
K and Ca use efficiency was found to be high at Pine
dominated stand. Statistical analysis revealed a
significant difference (p < 0.05) among the nutrient use
efficiency of the two species. The nutrient used
efficiency is inversely related to the availability of these
entire nutrients in soil. Since, it is indirectly act as an
indicator of soil nutrient availability while directly
related to the decomposition rate36. Both the litter
decomposition and the nutrient use efficiency are

Litter decomposition and nutrient cycling in forest of Kumaun region

Int. J. Fund. Appl. Sci. Vol. 4, No. 4 (2015) 91-98

Figure IV: (a) Nutrient Use Efficiency of two different plant species and (b) Nutrient retranslocation efficiency of selected
plant species

Table I: Soil physico-chemical characteristics under Pine

and Oak plantation in the study area
Soil Characteristics
Pine stand
Oak stand
pH

5.6870.015

5.8670.031

TOC (%)

5.7300.080

7.0800.130

TN (%)

0.2320.009

0.5550.001

AN (%)

0.0650.005

0.0710.005

TP (mg g-1)

0.1740.004

0.4000.003

AP (mg g-1)

0.0090.001

0.0340.001

AK (mg g-1)

0.1500.001

0.2670.002

ACa (mg g-1)

1.0670.003

1.7700.003

C/N

24.721.26

12.750.28

influenced by various factors including litter

composition, soil physico-chemical and biological
properties. Numerous studies have suggested that
recalcitrant like lignin, phenolics, tannins etc are also the
chief composition of litterfall that suppresses the
decomposition rate37, 38. However, these parameters are
not examined in the present study and needs to be
investigated further.
3.4 Nutrient retranslocation and cycling
The retranslocation efficiency of nutrients for Pinus
roxburgii and Quercus leucotricophora was represented
in (Fig. IVb). A significant difference (p < 0.05) was
observed in nutrient retranslocation of (N, P, K and Ca)
among the two species. The retranslocation percent of N,
P, K, Ca were 29.27%, 40.97%, 40.26% and 21.62% in
Pine whereas in Oak the percentage of these nutrients
were 20.31%, 28.37%, 35.26% and 17.61%, respectively.
Several authors have also demonstrated thar different
tree species have differed in their nutrients
retranslocation39, 40. The efficient retranslocation of the
essential nutrients is the most significant characteristic of
the climax phase of any forest ecosystem41. The
retranslocation efficiency of phosphorous and potassium

was recorded high among both the species suggested the

higher remobilization of the phosphorous and potassium
in the temperate forest. Furthermore, the NRE % of all
the nutrients was found to be high in Pine species
indicates comparatively lesser nutrient supply and soil
nutrient deficient condition than that of Oak. However,
the degree of retranslocation depends on its physical
properties as well as their requirement by the plant. The
lower NRE% in Oak species represents the lower
proportion of nutrients returns to litterfall. The
retranslocation with the reduction in restoration of
nutrients by leaves litter, results in independence of the
ecosystem with respect to soil and also a possibility of
managing the available elements in timed way42. The
difference in nutrient retranslocation is governed by
several factors including plant growth, Plant species,
plant age, site characteristics etc43. Additionally, these
responses are also dependent on the eco-physiological
response of tree species with the site environment and
are the strong indicator of soil fertility44.
3.5 Soil nutrient status
The pH of both the stands was recorded acidic in nature.
The detailed description of soil nutrient status of both the
stands was presented in (table I, II). The nutrient status of
Oak stand is found to be significantly superior to Pine
stand. Significant differences (p < 0.05) were recorded
between the two stands except AN (p > 0.05). The C/N
ratio is comparatively high at Pine stand which is mainly
related to decomposition rate and pattern of nutrient
immobilization. The decrease in value reveals increase in
decomposition process. Soil Microbial biomass (Carbon,
nitrogen and phosphorous) and basal respiration was
higher at Oak stand as compared to Pine stand.
Furthermore, the Oak stand displayed higher enzymes
activity than Pine stand. There is significant difference
recorded among the values of all the enzymes between
the two stands. The higher microbial biomass and
enzymatic activities at Oak stand suggested the higher
availability of nutrient at Oak stand than that of Pine
stand. The plants grow within nutrient-poor soil
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Chandra et al.

Int. J. Fund. Appl. Sci. Vol. 4, No. 4 (2015) 91-98

Table II: Soil microbial characteristics enzymatic activities

under Pine and Oak plantation in the study area
Microbial
Characteristics

Pine Stand

Oak Stand

MBC(g g-1)

335.20 2.42

448.22 2.31

MBN(g g-1)

72.80 1.51

91.38 2.18

MBP(g g-1)

34.47 1.60

62.17 1.54

Basal respiration
(g CO2 g-1 soil hr-1)

23.22 0.21

25.54 0.21

Enzymes Assay
Acid Phosphates
-Glucosidase
DHA
Protease
Cellulases

2287.70 44.64
586.34 20.52
224.79 13.53
3702.48 39.00
0.40 0.02

3235 121.70
1621.78 70.03
623.09 8.88
4022.73 41.00
0.73 0.03

FDA

356.05 6.63

491.77 24.15

MB(C), MB(N), MB(P)-Microbial Biomass (Carbon), (Nitrogen),

(Phosphorous)., DHA-Dehydrogenase., FDA-Fluorescein diacetate.

displayed high nutrient resorption resultant to low litter

quality and decomposition rates45 that leads to the lower
magnitude of nutrient return to soil and litter. It has been
observed that broad leaved tree may change the forest
microenvironment in order to have faster decomposition
and this was in favor of oak dominated stand where
microbial activity is high46, 35.

4. Conclusion
The study emphasizes that the two plant species differ in
litterfall plants displayed different rate of decomposition.
Also, the retranslocation of nutrients influenced not only
by litter decomposition but also by the soil properties as
well as plant nutrient status. It has been concluded that
the high enzymatic activity and microbial biomass at Oak
stand signifies that the availability of nutrients was
higher at Oak stand than the Pine stand. Additionally, the
Pine stand was undersupplied nutrient ecosystem
characterized as high nutrient use efficiency and retranslocation of nutrients, lead to the depletion in litter
quality as well as decomposition rate on the floor of Pine
stand. Furthermore, the nutrient status of leaves, litter
and soil play equal role in the working of nutrient cycle
and the survival of forest ecosystem.

Acknowledgments
We are grateful to director CSIR-NBRI, Lucknow for
providing necessary facilities and encouragement.
Financial support received from UGC during the
research is duly acknowledged. This work was supported
by CSIR-Network Project BSC-0109 (SIMPLE) and by
DST (Grant No.DST/CCP/PR/03/2011).

Conflict of interest
The authors declares none.

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