Respiratory costs of woody tissues in a Quercus pyrenaica coppice - SISEF.ORG
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iForest Short Communication
doi: 10.3832/ifor2599-011
vol. 11, pp. 437-441
Biogeosciences and Forestry
Respiratory costs of woody tissues in a Quercus pyrenaica coppice
Roberto Luis Salomón (1-2), Long-term coppicing leads to the development of massive root systems. A dis-
proportionate carbon investment in root maintenance has been pointed as a
Jesús Rodríguez-Calcerrada (1), cause of the widespread decline of abandoned coppices. We aimed at assessing
Luis Gil (1), how coppicing has influenced root and shoot development and related carbon
María Valbuena-Carabaña (1) loss ascribed to maintenance of woody tissues in Quercus pyrenaica. For this
goal, results from published studies on root dynamics, woody biomass and
respired CO2 fluxes in an abandoned Q. pyrenaica coppice were integrated
and extended to quantify overall respiratory expenditures of above- and be-
low-ground woody organs. Internal and external CO 2 fluxes together with soil
CO2 efflux were monitored in eight stems from one clone across a growing sea-
son. Stems and roots were later harvested to quantify the functional biomass
and scale up root and stem respiration (RR and RS, respectively) to the clone
and stand levels. Below- and above-ground biomass was roughly equal. How-
ever, the root-to-shoot ratio of respiration (RR/RS) was generally below one.
Relatively higher RS suggests enhanced metabolic activity aboveground during
the growing season, and highlights an unexpected but substantial contribution
of RS to respiratory carbon losses. Moreover, soil and stem CO 2 efflux to the at-
mosphere in Q. pyrenaica fell in the upper range of reported rates for various
forest stands distributed worldwide. We conclude that both RS and RR repre-
sent an important carbon sink in this Q. pyrenaica abandoned coppice. Com-
paratively high energetic costs in maintaining multiple stems per tree and cen-
tennial root systems might constrain aboveground performance and contribute
to coppice stagnation.
Keywords: Carbon Loss, CO2 Fluxes, Coppice Stagnation, Oak, Resprouting
Species, Root Respiration, Stem Respiration
Introduction and consequently, hindered application of coppicing have been suggested as a poten-
Quercus pyrenaica Willd. is a vigorous alternative management plans. Due to the tial driver of Q. pyrenaica decay (Bravo et
root-resprouting species that has been in- wide distribution and significant ecological al. 2008), but assessments on carbon ex-
tensively coppiced for firewood, charcoal value of coppiced stands of Mediterranean penditures have not been essayed to date.
and woody pastures for centuries (Ruiz de oak species, silviculture faces the crucial Quantification of the relative weight of res-
la Torre 2006). Due to the appearance of challenge of finding new alternative uses piratory carbon sinks for the plant is crucial
new energy sources and rural exodus that for these abandoned coppices (Cañellas et for a better understanding of tree carbon
occurred in the 1970s, coppicing has mostly al. 2004, Bravo et al. 2008). Attempts to budgets (Waring et al. 1998, Amthor 2000,
ceased, and symptoms of decline – slow conversion into high forests via thinning Rambal et al. 2014). Notwithstanding, our
stem growth, branch dieback, and scarce have not been successful to date, partly comprehension of respiratory processes,
acorn production – are widely observed in due to the lack of a comprehensive under- particularly of woody organs, is limited
abandoned stands (Serrada & Bravo 2012). standing of the physiological mechanisms compared to our knowledge of photosyn-
Coppice stagnation entails ecologic, eco- underpinning tree stagnation (Salomón et thesis (Guidolotti et al. 2013, Rambal et al.
nomic and social problems, namely in- al. 2017). 2014, Huntingford et al. 2017). In resprout-
creased fire risk, stand over-aging, low pro- Disproportionate respiratory costs of ing deciduous species, nonstructural carbo-
ductivity, absence of sexual regeneration, large root systems grown after centennial hydrates are stored in large amounts in
woody organs (Bond & Midgley 2001) that
can contain a large portion of living paren-
(1) Forest Genetics and Ecophysiology Research Group, E.T.S. Forestry Engineering, Tech- chyma (Rodríguez-Calcerrada et al. 2015).
nical University of Madrid, Ciudad Universitaria s/n, 28040, Madrid (Spain); (2) Laboratory of The penalty in terms of respiratory carbon
Plant Ecology, Department of Applied Ecology and Environmental Biology, Faculty of Bio- loss associated to the maintenance of
science Engineering, Ghent University, Coupure links 653-9000 Ghent (Belgium) these storage tissues (Landhäusser & Lief-
fers 2002, Drake et al. 2009) could be of
@ Roberto Luis Salomón (robertoluis.salomonmoreno@ugent.be) particular relevance in carbon budgets of
root-resprouting Q. pyrenaica. Long lasting
Received: Aug 11, 2017 - Accepted: Apr 09, 2018 coppicing might lead to massive systems of
living roots (Salomón et al. 2016a, Vrška et
Citation: Salomón RL, Rodríguez-Calcerrada J, Gil L, Valbuena-Carabaña M (2018). al. 2016) that store but also consume a
Respiratory costs of woody tissues in a Quercus pyrenaica coppice. iForest 11: 437-441. – doi: large portion of carbohydrates assimilated
10.3832/ifor2599-011 [online 2018-06-18] aboveground.
To better understand the role of respira-
Communicated by: Giorgio Matteucci tory carbon loss in Q. pyrenaica decay, we
gathered and extended previous published
© SISEF http://iforest.sisef.org/ 437 iForest 11: 437-441
Salomón RL et al. - iForest 11: 437-441
work on Q. pyrenaica biomass and respira- na-Carabaña & Gil 2013 – see Fig. 1 in Sa- from McGuire & Teskey 2004). FT was calcu-
iForest – Biogeosciences and Forestry
tion of woody tissues (Salomón et al. 2015, lomón et al. 2015). Data on woody tissue lated as a function of the sap flux and the
2016a, 2016b, 2016c) to scale up stem and respiration was collected from the eight vertical gradient of CO 2 dissolved in sap so-
root respiration (RS and RR, respectively) to stems belonging to a single clonal geno- lution (sap [CO2*]). Sap flux density was
the tree and stand levels. We aimed at type (clone). Four 24-h measurement cam- measured using Granier-type thermal dissi-
comparing respiratory expenditures (i) be- paigns were conducted across the growing pation probes, and sap [CO 2*] was esti-
tween above- and below-ground woody or- season of 2013, at the end of which stems mated from measurements of xylem [CO 2]
gans across one growing season, and (ii) in and roots were harvested for biomass in the gas phase, sap temperature, and sap
relation to data gathered from various quantification and scaling up of RS and RR to pH in each stem applying Henry’s law.
forests to provide an insight of the magni- the clone and stand levels. The root system Briefly, xylem [CO2] was measured with
tude of carbon invested for woody tissue was hydraulically excavated with a high- solid non-dispersive infrared (NDIR) CO 2
respiration in Q. pyrenaica coppices. We ex- pressure water pump down to 1 m depth sensors (model GMM221®, Vaisala, Helsinki,
pect RS and particularly RR to be important over an area of 81 m2 (Fig. 1). Biomass was Finland) inserted into the stem above and
carbon sinks for the plant (i.e., from the partitioned into leaves, branches, stems, below the stem collar. Stem temperature
plant perspective), and therefore high taproots, coarse roots and fine roots. was measured with type-T thermocouples
RR/RS ratios as well as woody tissue respira- Woody biomass was further partitioned inserted 5 cm away from the NDIR probe.
tion rates relative to other forest stands. into bark, sapwood and heartwood tissues Sap pH was measured with a micro-pH
from allometric equations adjusted by electrode and a portable pH meter (Crison,
Materials and methods means of exhaustive sampling of branches, Barcelona, Spain) on sap samples ex-
To estimate respiratory carbon loss of stems, taproots and coarse roots. Leaf pressed from detached twigs using a pres-
woody organs in a coppice system of Q. area index (LAI) was estimated from mea- sure chamber (see Salomón et al. 2016b for
pyrenaica, we reviewed our previous work surements of specific leaf area of sampled further details). Overall EA, FT and RS at the
on Q. pyrenaica root development and leaves and total leaf biomass. Further de- clone level were estimated by aggregating
biomass (Salomón et al. 2016a), and inter- tails on stand characteristics, excavation EA, FT and RS scaled up for each stem (and
nal and external stem CO2 fluxes (Salomón methodology, and above- and below- their branches) according to their corre-
et al. 2015, 2016b, 2016c), together with un- ground biomass measurements can be sponding volume of living woody biomass.
published data of soil CO 2 efflux. These seen in Salomón et al. (2016a). Aboveground clonal respiratory fluxes
studies were performed in a one-hectare Stem CO2 efflux to the atmosphere (EA) were eventually expressed on a soil surface
experimental plot located in the Monte was measured in every stem with a porta- area basis taking into account the clonal
Matas de Valsaín (Segovia, Spain) at an alti- ble infrared gas analyzer (LI-6400®, Li-Cor surface extension.
tude of 1140 m a.s.l. Climate is sub-Mediter- Inc., Lincoln, NE, USA) and a soil chamber Soil CO2 efflux (ES) was measured with a
ranean with an average annual rainfall and (LI-6400-09®) using PVC collars attached to portable infrared gas analyzer and a soil
temperature of 885 mm and 10 °C, respec- the stems. Stem EA measured on a surface chamber using soil PVC collars (see Sa-
tively. It consists on a monospecific one- area basis (EA(S)) was expressed on a vol- lomón et al. 2015 for detailed methodol-
storied regular coppice of Q. pyrenaica with ume basis (EA(V) – eqn. 1): ogy). Unpublished data from four soil col-
a stand density of 781 stems ha -1. The forest S lars located below the canopy of the eight
E A( V )= E A(S) =
has been subjected to coppicing since at V monitored stems were averaged to obtain
least the XII century, and traditional man- 2 π r h+l L 2r (1) clonal ES on a soil area basis (m 2). Since
agement was abandoned in the 1970s. = E A(S) =E A( S) 2 h+ l 2 roots barely extended beyond the exca-
π (r2h+ l−r 2h ) L (r h+l −r h )
Stems within the plot were geo-referenced vated area (81 m2), a buffer of 0.63 m was
and leaves collected for genetic analyses to where S and V are the axial surface area added to estimate the clone extension (102
delineate the commonly inconspicuous and the volume of living tissues (bark and m2). This buffer distance was chosen to
clonal structure of Q. pyrenaica coppiced sapwood) of the stem segment, respec- meet actual stand density (8 stems in 102
stands. Note that Q. pyrenaica is a root re- tively; rh and rh+l denote the radius of heart- m2 = 784 stems ha-1) and scale up results to
sprouting species, hence one clone can be wood and heartwood plus living tissues, re- the stand level. Root-respired CO 2 diffusing
constituted by several stems located far spectively, and L is the vertical length of to the atmosphere through soil (ES-ROOT)
away (dozens of meters) from each other the stem segment (Salomón et al. 2016c). was estimated from ES measurements and
(Valbuena-Carabaña & Gil 2017). The clonal Stem respiration (RS) was estimated as the the relative contribution of autotrophic
assignment of stems was based on nuclear sum of EA(V) and the internal CO2 flux respiration to ES. Seasonality of root au-
microsatellite molecular markers (Valbue- through xylem (FT) as RS = EA + FT (adapted totrophic contribution to ES was obtained
Fig. 1 - Root systems of
two hydraulically exca-
vated clones of Quercus
pyrenaica located in the
Monte Matas de Valsaín
(Segovia, Spain). The root
system of the large clone
surveyed in this study
(lower part of the photo-
graph) covered at least 81
m2. Before harvesting, the
clone was composed by
eight stems connected
through root grafts and
parental roots.
438 iForest 11: 437-441Woody tissue respiration in oak coppice
from two studies in a Mediterranean Quer-
iForest – Biogeosciences and Forestry
cus cerris coppice cut one (Rey et al. 2002)
and 17 (Tedeschi et al. 2006) years before
ES measurements. Spring contributions re-
ported in these studies were attributed to
the first measurement campaign (DOY 143-
144), summer contributions to the second
(DOY 183-184) and third (DOY 218-219) cam-
paigns, and autumn contributions to the
fourth campaign (DOY 266-267). To ac-
count for FT on RR estimates (RR = ES-ROOT + FT
– Aubrey & Teskey 2009), internal and ex-
ternal CO2 fluxes were measured at the
base of the stem (0.1 m).
To compare ES, RR and RS averaged over
the growing season within the monitored
clone, ANOVA and pairwise comparisons
were performed in R software (version
3.4.0). Respiratory fluxes at the stand level
from 15 sites were gathered (Tab. S1 in Sup-
plementary material) to evaluate the rela-
tive magnitude of respiratory costs of the
Fig. 2 - Diel variations in stem CO2 efflux to the atmosphere (EA), stem internal CO2
surveyed Q. pyrenaica coppice. Inter-site
transport through xylem (FT) and soil CO2 efflux (ES) on four dates over the growing
statistical comparisons were not per-
season in an abandoned coppice of Quercus pyrenaica. Fluxes registered from one
formed because only one site was sur-
stem segment and one soil collar intensively monitored (18 times day -1) are shown.
veyed in this study.
Additional stem segments and collars used to average EA, FT and ES are not displayed in
this figure as they were monitored less intensively (4 times day -1). EA and FT on a vol-
Results and discussion
ume basis was obtained from previous work (Salomón et al. 2016c) and expressed on
Seasonal and diel variation in EA, FT and ES
a soil area basis for comparison with ES. Shaded areas indicate night-time.
on a soil area basis are shown in Fig. 2. The
contribution of FT to aboveground RS was
less than 10% (Salomón et al. 2016c), where- aboveground and 972 Kg belowground (Sa- one along the growing season evidenced
as FT belowground was less than 2% of lomón et al. 2016a). Consequently, sea- an unexpected large weight of above-
ES-ROOT (Salomón et al. 2015). The modest sonal deviations of RR/RS from unity re- ground woody tissue respiration as a car-
contribution of axial CO2 transport to total flected differences in the metabolic activity bon sink for the plant. The accumulation of
respiration rates is explained by the low between below- and above-ground organs woody biomass in stems and branches in
xylem [CO2] observed in Q. pyrenaica, gen- over time. RR/RS ratio above one was this over-aged coppice (cut for the last
erally lower than 1%. This concentration is uniquely observed during spring (Tab. 1), time around 1967), together with the re-
about one order of magnitude lower than likely explained by an earlier growth of markably high portion of living paren-
that reported for other tree species using roots relative to stems (López et al. 2001), chyma observed in Q. pyrenaica stems (Ro-
this methodology (Teskey et al. 2008). The particularly by intense fine root growth dríguez-Calcerrada et al. 2015) may contrib-
limited build-up of CO2 in the xylem was and belowground cambial activity at this ute to the high respiratory costs of woody
partly ascribed to the low resistance to ra- time of the year (Courty et al. 2007). The organs aboveground.
dial CO2 diffusion, likely related to the poor decrease in RR/RS observed onward (ratios Respiratory fluxes at the stand level, and
plant water status of species distributed below one) resulted from the moderate extrapolated to the whole year, were com-
across drought-prone regions (Salomón et root activity relative to the intensification pared with those reported for several for-
al. 2016b). Averaged over the growing sea- of aboveground metabolism, namely stem est sites (see Tab. s1 in Supplementary ma-
son, ES (38.9 mol CO2 clone-1 day-1) was growth, leaf development and phloem terial for details on the extrapolation). Av-
greater than RS (11.7 mol CO2 clone-1 day-1) transport. Predominant RR/RS ratios below erage ES and EA across 15 stands were 776
and RR (8.5 mol CO2 clone-1 day-1 – P < 0.001;
Tab. 1, Fig. 2), being RS and RR not signifi-
cantly different (P > 0.10). Due to the mag- Tab. 1 - Above- and below-ground respiratory fluxes in an eight-stemmed clone of
nitude of ES, the contribution of auto- Quercus pyrenaica. Stem CO2 efflux to the atmosphere (EA), soil CO2 efflux (ES), and
trophic respiration to ES substantially deter- stem and root respiration (RS and RR, respectively) were measured during four 24-h
mined the root-to-shoot ratio of respira- campaigns throughout 2013 growing season. (a): RS was estimated as the sum of EA
tion (RR/RS). To illustrate this, a contribution and the internal CO2 flux through xylem (FT). (b): RR was estimated from ES measure-
of RR to ES ranging between 14 and 27% ments and the contribution of autotrophic respiration to ES. Autotrophic contribution
(Rey et al. 2002, Tedeschi et al. 2006 – data to ES was obtained from two reports of a Mediterranean coppice of Quercus cerris cut
reported for a Mediterranean oak coppice one (Rey et al. 2002) and 17 years (Tedeschi et al. 2006) before measurements. Esti-
at different states of maturity) yielded mations of RR and RR/RS ratios from contributions reported in both studies (recently
RR/RS ratios ranging between 0.42 and 1.18 coppiced vs mature stand) are shown in left and right sub-columns, respectively. FT
across the growing season (Tab. 1). Alter- was neglected in RR due to its low contribution (< 2%).
natively, if heterotrophic and autotrophic
contributions to ES were considered equal, Campaign EA RS (a) ES RR (b)
RR/RS
as generally assumed for different forest DOY (mol CO2 clone-1 day-1)
biomes (Hanson et al. 2000, Aubrey & 143-144 6.16 6.16 26.83 5.03 vs 7.25 0.82 vs 1.18
Teskey 2009), RR/RS would reach values 183-184 12.28 13.13 52.14 11.32 vs 11.70 0.86 vs 0.89
ranging from 1.36 to 2.18. 218-219 15.63 16.45 43.72 9.49 vs 9.82 0.58 vs 0.60
Above- and below-ground functional 266-267 10.54 11.23 32.76 8.82 vs 4.68 0.79 vs 0.42
woody biomass (bark and sapwood) was
Mean 11.15 11.74 38.86 8.66 vs 8.36 0.76 vs 0.77
similar in the surveyed clone: 1026 Kg
iForest 11: 437-441 439Salomón RL et al. - iForest 11: 437-441
and 162 g C m-2 year-1, respectively, whereas Acknowledgements
iForest – Biogeosciences and Forestry
ables and stand structure on ecosystem respi-
these values increased up to 1164.2 and We are grateful to Javier Donés (Centro ration components in a Mediterranean beech
297.0 g C m-2 year-1 in the surveyed Q. pyre- de Montes y Aserradero de Valsaín) for eco- forest. Tree Physiology 33: 960-972. - doi:
naica coppice (Tab. S1). That is, ES and EA nomic and logistical support. We also thank 10.1093/treephys/tpt065
were 1.5 and 1.8 times higher in this study, Elena Zafra, Guillermo González, César Hanson PJ, Edwards NT, Garten CT, Andrews JA
suggesting greater carbon losses from soil Otero, Manuel Iglesias, Paula Guzmán, Aida (2000). Separating root and soil microbial con-
and stem respiration. These ratios could be Rodríguez, Jose Carlos Miranda, Rosa Ana tributions to soil respiration: A review of meth-
substantially reduced, however, if xylem López, Eva Miranda and Matías Millerón ods and observations. Biogeochemistry 48: 115-
transport of respired CO2 was accounted for their enthusiastic help with field work. 146. - doi: 10.1023/A:1006244819642
for (Aubrey & Teskey 2009) in the 15 forest This work was funded by the Comunidad Hernández-Santana V, Martínez-Vilalta J, Martí-
stands. Leaf area index, a key ecophysio- de Madrid through CAM P2009/AMB-1668 nez-Fernández J, Williams M (2009). Evaluating
logical determinant of carbon gas ex- and S2013/MAE-2760 projects and by the the effect of drier and warmer conditions on
change at the canopy (Bréda 2003), was Organismo Autónomo de Parques Nacio- water use by Quercus pyrenaica. Forest Ecology
additionally considered for comparison. nales through the PPNN 1148/2014 project. and Management 258: 1719-1730. - doi: 10.1016/j.
LAI in our site was 3.8, a value that falls Roberto L. Salomón was supported by a foreco.2009.07.038
within the reported range for the species Ph.D. scholarship from the Universidad Huntingford C, Atkin OK, Martinez de la Torre A,
(Hernández-Santana et al. 2009) and the Politécnica de Madrid. Mercado LM, Heskel MA, Harper AB, Bloom-
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