Respiratory costs of woody tissues in a Quercus pyrenaica coppice - SISEF.ORG

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Respiratory costs of woody tissues in a Quercus pyrenaica coppice - SISEF.ORG
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
Respiratory costs of woody tissues in a Quercus pyrenaica coppice - SISEF.ORG
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: 437-441
Woody 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                                                                                                                          439
Salomón 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-
                                        genus Quercus (Bréda 2003). The slightly                                                                  field KJ, Sullivan OS, Reich PB, Whythers KR,
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