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Applied Soil Ecology
journal homepage: www.elsevier.com/locate/apsoil
Foliage C:N ratio, stage of organic matter decomposition and interaction
with soil affect microbial respiration and its response to C and N addition
more than C:N changes during decomposition
Veronika Jílková
a
, Petra Straková
b,c
, Jan Frouz
a,d,
⁎
a
Biology Centre, Czech Academy of Sciences, Institute of Soil Biology and SoWa RI, Na Sádkách 7, České Budějovice CZ-37005, Czech Republic
b
Department of Forest Sciences, Peatland Ecology Group, University of Helsinki, Latokartanonkaari 7, Helsinki FI-00014, Finland
c
Natural Resources Institute Finland (Luke), Latokartanonkaari 9, Helsinki FI-00790, Finland
d
Charles University in Prague, Institute for Environmental Studies, Benátská 2, Prague CZ-12801, Czech Republic
ARTICLE INFO
Keywords:
Light fraction
Heavy fraction
Litter
Nutrients
Carbon dioxide
Spoil heaps
ABSTRACT
How litter at various stages of decomposition reacts to C and N additions is unclear. Here we used five substrates
(litter, fermentation [Oe] layer, bulk soil, and the light fraction [LF] and heavy fraction [HF] of SOM) obtained
from sites supporting five plant monocultures (Alnus glutinosa, Quercus robur, Salix caprea, Calamagrostis epigejos,
or Picea omorica) with foliage C:N ratios ranging from 17 to 48. These plant-specific communities were ex-
perimentally planted on a post-mining heap and had affected the substrates used in this study for 40 years. Soils
and other environmental factors were similar among the sites. Substrates were incubated for 3 weeks without
nutrient addition or with C (glucose) or N (ammonium nitrate) addition, and microbial respiration was de-
termined weekly. Substrate C:N ratios were determined at the start of the incubation and were highest for litter
followed by Oe layer > LF > bulk soil and HF. Foliage C:N ratio was a better indicator of microbial respiration
than the substrate C:N ratio, suggesting that the foliage C:N ratio reflected unmeasured leaf properties that
determined microbial respiration. Respiration was highest in the litter followed by Oe layer > bulk
soil > LF > HF. C addition increased respiration of the bulk soil (+39%), LF (+48%), and HF (+72%).
Priming of SOM respiration was therefore higher in substrates with less available C. N significantly increased
respiration of litter (+19%) but decreased respiration of bulk soil (-18%). The difference in respiration of HF
vs. bulk soil following N addition suggested that, in addition to the stage of decomposition, environmental
properties present in bulk soil but absent in HF may cause the reduction in respiration after N addition to bulk
soil. Overall, the results indicate that differences in the contents of SOM fractions among soils will affect the
responses of those soils to C and N additions.
1. Introduction
The carbon dioxide (CO
2
) concentration in the atmosphere has in-
creased to almost 400 ppm (IPCC, 2014), and higher carbon (C) inputs
to soils are thus expected due to increased litter production and rhi-
zodeposition (Cotrufo et al., 1998a, 1998b; Coûteaux et al., 1999;
Norby et al., 2005; Phillips et al., 2006). Free-air CO
2
enrichment stu-
dies corroborate these expectations in that elevated CO
2
(ambient
+200 ppm) causes significant increases in litterfall mass and/or root
increment (e.g., Allen et al., 2000; Hoosbeek and Scarascia-Mugnozza,
2009; Lukac et al., 2009). Moreover, most ecosystems are experiencing
increased inputs of anthropogenically derived nitrogen (N) (Galloway
et al., 2008; Schlesinger, 2009; Vitousek et al., 1997). Several studies
show that a large-scale N input to soil by enhanced fertilization and
atmospheric deposition is likely to reduce soil respiration (Janssens
et al., 2010; Schlesinger and Andrews, 2000). When sites are not N-
limited, a long-term N addition also decreases soil respiration (Guo
et al., 2017), but here mechanisms are likely to be more complex and
include soil acidification and a shift in microbial community composi-
tion. Because C storage or content is nearly three times greater in soil
than in aboveground biomass and is approximately two times greater
than in the atmosphere (Eswaran et al., 1993; Schlesinger, 1977), an
important question is “How will soils react to such high C and N in-
puts?” An understanding of the response of soil organic C (SOC) to
https://doi.org/10.1016/j.apsoil.2020.103568
Received 23 October 2019; Received in revised form 18 February 2020; Accepted 20 February 2020
⁎
Corresponding author at: Biology Centre, Czech Academy of Sciences, Institute of Soil Biology and SoWa RI, Na Sádkách 7, České Budějovice CZ-37005, Czech
Republic.
E-mail address: frouz@natur.cuni.cz (J. Frouz).
Applied Soil Ecology 152 (2020) 103568
0929-1393/ © 2020 Elsevier B.V. All rights reserved.
T