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Correspondence Hemicelluloses account for one-quarter of the global dry plant biomass
*Corresponding author, and therefore are the second most abundant biomass fraction after
e-mail: c.schaedel@unibas.ch
cellulose. Despite their quantitative significance, the responsiveness of
Received 15 November 2009; hemicelluloses to atmospheric carbon oversupply is still largely unknown,
revised 20 January 2010 although hemicelluloses could serve as carbon sinks with increasing CO2
concentrations. This study aimed at clarifying the role hemicelluloses
doi:10.1111/j.1399-3054.2010.01360.x play as carbon sinks, analogous to non-structural carbohydrates (NSC), by
experimentally manipulating the plants’ carbon supply. Sixteen plant species
from four different plant functional types (grasses, herbs, seedlings of broad-
leaved trees and conifers) were grown for 2 months in greenhouses at either
extremely low (140 ppm), medium (280 ppm) or high (560 ppm) atmospheric
CO2 concentrations, thus inducing situations of massive C-limitation or
-oversupply. Above and belowground biomass as well as NSC significantly
increased in all species and tissues with increasing CO2 concentrations.
Increasing CO2 concentrations had no significant effect on total hemicellulose
concentrations in leaves and woody tissues in all species, except for two out
of four grass species, where hemicellulose concentrations increased with
atmospheric CO2 supply. Despite the overall stable total hemicellulose
concentrations, the monosaccharide spectra of hemicelluloses showed a
significant increase in glucose monomers in leaves of woody species as
C-supply increased. In summary, total hemicellulose concentrations in de
novo built biomass seem to be largely unaffected by changed atmospheric
CO2 concentrations, while significant increases of hemicellulose-derived
glucose with increasing CO2 concentrations in leaves of broad-leaved and
conifer tree seedlings showed differential responses among the different
hemicellulose classes in response to varying CO2 concentrations.
Abbreviations – DM, dry matter; HPLC, high performance liquid chromatography; HSD, honestly significant difference; NSC,
non-structural carbohydrates.
(Alopecurus and Arrhenaterum) in which increasing CO2 significant effect of CO2 on sapwood hemicellulose
concentration significantly increased total hemicellulose concentrations in any of the eight investigated tree
concentration by about 14% (Fig. 4A). Although Silene species. Likewise, hemicellulose concentrations in roots
showed a 46% increase of hemicellulose concentrations did not show any CO2 effect (data not shown).
from 140 to 560 ppm CO2 , this increase was not Because the strong increase of NSC with CO2
significant because of the high variability between concentrations in most species might mask (‘dilute’)
individuals. Hemicellulose concentrations in seedlings more moderate increases of other cell compounds,
of broad-leaved trees and conifers were constant over hemicelluloses were additionally calculated not only
all CO2 treatments except in Tilia where 34% more relative to the total dry weight but also on an
hemicelluloses were measured at 140 ppm than at NSC-free basis. The general pattern of hemicellulose
the two higher CO2 concentrations. Averaged over all concentrations on an NSC-free basis in leaves and
species of one plant functional type, no CO2 effect was sapwood in response to increasing CO2 concentrations
found for the total hemicellulose concentrations (Fig. 4B, did not differ from that on a % DM basis, but became
Table 3). more pronounced, especially with an average 20% (not
Hemicellulose concentrations in sapwood were significant) increase in the hemicellulose concentration
similar for seedlings of broad-leaved trees and conifers in grass leaves and a 17% (not significant) increase in
and ranged from 15 to 23% with slightly higher values needles of conifer seedlings from 140 to 560 ppm CO2
in broad-leaved tree seedlings (Fig. 5). There was no (data not shown).
Table 3. ANOVA table of the fixed effects from the mixed effects model plant functional group (Figs 6 and 7). Hemicelluloses of
analysis for the increase in concentrations of NSC and hemicelluloses in
grass leaves contained over 60% xylose, c. 20% arabi-
leaves and sapwood of four different plant functional groups. Broad-
leaved trees and conifers produced sapwood. P values smaller than 0.05
nose and mainly galactose, glucose and fucose within
are in bold the remaining 20%. Xylose, arabinose and galactose
were the major sugars in leaves of herbaceous species
NSC Hemicelluloses with smaller (8–10%) proportions of rhamnose and glu-
df F P df F P cose. Leaves of broad-leaved tree seedlings revealed a
Leaves high proportion of xylose, likely deriving from xylans,
CO2 2 38.7 <0.03 2 0.19 0.84 while mannose was the major monosaccharide in nee-
Functional group 3 7.9 <0.001 3 11.13 <0.001 dles of conifer seedlings, indicating high proportions of
CO2 × functional group 6 1.0 0.40 6 0.25 0.96 mannans with also arabinose, galactose and glucose as
Sapwood
sugar residues (Fig. 6). In leaves of broad-leaved tree
CO2 2 2.6 0.28 2 0.02 0.98
Functional group 1 514.6 <0.001 1 66.6 <0.001
seedlings and conifer seedlings significantly (P < 0.04
CO2 × functional group 2 0.3 0.76 2 0.78 0.46 by linear mixed effects model) higher glucose concentra-
tions were measured with increasing CO2 concentrations
(Fig. 6). Although present in leaves of all tree species, the
increase in glucose content was especially pronounced
Monosaccharide spectrum of hemicelluloses
in Acer, Quercus, Larix and Picea. However, because of
The analysis of the monosaccharide spectra of hemi- the relatively small proportion of glucose within the total
celluloses by HPLC revealed specific patterns for each hemicelluloses, this increase in glucose concentration
Total hemicelluloses
Discussion
Unlike NSC, the overall response of total hemicellulose
Biomass and NSC responses to elevated CO2
concentrations to changes in the carbon supply was very
concentrations
weak for all tissues and species in the current study.
The successful manipulation of the plants’ carbon supply Only two out of four grass species (Alopecurus and
was evident by the significant changes in growth Arrhenaterum) responded significantly to elevated CO2
concentrations with 14% higher hemicellulose concen- among the studies. Peñuelas et al. (2002) found increas-
tration in leaves at 560 ppm compared with 140 ppm ing hemicellulose concentrations with increasing CO2
CO2 . All other species revealed constant hemicellu- concentrations in leaves of the Mediterranean shrub
lose concentrations with increasing CO2 concentrations, Myrtis communis, whereas two other shrub species (Erica
except Tilia which was the only species that unexpect- arborea and Juniperus communis) growing at the same
edly showed significantly higher hemicellulose concen- site showed no change in hemicellulose concentration
trations in leaves developed at 140 ppm compared with to elevated CO2 . The authors explained this inconsis-
higher CO2 concentrations; a phenomenon which needs tency with different carbon investment strategies among
further explanation. The overall picture of stable hemi- different plant species. Within another study, leaf lit-
cellulose concentrations prevailed, even if the ‘diluting’ ter of Castanea sativa saplings grown at 700 ppm CO2
effect of the marked changes in NSC was considered revealed 23% higher hemicellulose concentrations than
and hemicellulose concentrations were expressed on a at ambient conditions (Couteaux et al. 1996). Similarly,
% NSC-free DM basis. Hall et al. (2006) found increased hemicellulose concen-
Hemicelluloses have rarely been studied in relation trations with elevated CO2 concentrations in leaf litter of
to increasing CO2 concentrations. Most of those stud- Quercus myrtifolia. Leaves of Betula pendula revealed
ies were restricted to woody species and the reported significantly higher hemicellulose concentrations under
effects were rather inconsistent. In addition, the focus elevated CO2 but this effect disappeared again when
of these studies had not been on the reaction of hemi- leaves were exposed to ozone (Oksanen et al. 2005).
celluloses, and the quality and precision of the different No change in hemicellulose concentration at increased
methods used to determine hemicelluloses varied greatly CO2 concentrations was found in wood of 15-year old
hemicelluloses has not been investigated so far. or litter decomposition (Couteaux et al. 1996). Within
A significant change of the monosaccharide composition this study, glucose was the only monosaccharide that
of hemicelluloses, as well as a pronounced change showed significantly higher proportions with increasing
of hemicellulose concentrations in plant tissue, could CO2 concentrations in most of the investigated
ultimately affect ecosystem processes like herbivory tissues, which led to significantly higher proportions
Conclusions
This study is the first broad-scale investigation on cell
wall hemicelluloses in response to varying CO2 concen-
trations in different plant functional types. Only leaves
of two grass species exhibited a significant increase
in hemicellulose concentration with increasing CO2
concentrations, while all other plant functional types
did not show significant changes in the hemicellulose
concentration with C-source–sink imbalances. Thus,
hemicellulose concentrations in de novo built biomass
seem to be largely unaffected by changed atmospheric
CO2 concentrations. In contrast, significant increases
of hemicellulose-derived glucose with increasing CO2
concentrations in leaves of broad-leaved and conifer
tree seedlings showed that hemicellulose compositions
change in response to varying CO2 concentrations.
Overall, this experiment showed that the structural
functions of hemicelluloses do not generally allow for
pronounced variations of the total hemicellulose con-
tent of tissue built under situations of different carbon
availabilities. In conclusion, a quantitative increase of
cell wall hemicelluloses in plants, similar to this study,
at future high atmospheric CO2 concentrations is not
expected.
Fig. 7. Fractions of hemicellulose-derived monosaccharides in sapwood
of all species from broad-leaved trees and conifers determined by HPLC.
Acknowledgements – This work was supported by the
Values are means of two chamber means.
Swiss National Science foundation project no. 3100A0-
17548. The authors would like to thank Georges Grun
of glucose in the total hemicellulose pools of tree for setup and maintenance of the CO2 controlling system,
leaves. The strong responsiveness of glucose toward Olivier Bignucolo for NSC analyses and Pascal Niklaus for
changed CO2 supplies can be most likely explained statistical help.
Edited by B. Sundberg