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Applied Clay Science
journal homepage: www.elsevier.com/locate/clay
Research paper
Carbohydrate concentrations and enzyme activities as influenced by
exchangeable cations, mineralogy and clay content
Fatemeh Rakhsh
⁎
, Ahmad Golchin
Soil Science Department, Faculty of Agriculture, University of Zanjan, Zanjan, Iran.
ARTICLE INFO
Keywords:
Acid phosphatase
Alfalfa residues
Alkaline phosphatase
Carbohydrate
CM-cellulase
Homoionic clay
ABSTRACT
Enzymes can exist in bound or free form within the soil, but the impact of interactions between extracellular
enzymes with the amount and type of clay minerals and exchangeable cations on the activities of enzymes and
decomposition or retention of carbohydrates in soil is still poorly understood. Appropriate amounts of homoionic
Na-, Ca- and Al-clay minerals from Georgia kaolinite, Illinois illite and Wyoming montmorillonite were mixed
with pure sand to prepare artificial soils with different clay contents, exchange cations and clay types to examine
the effects of exchangeable cations, mineralogy and clay content on the concentrations of hot water and dilute
acid extractable carbohydrates and activities of acid and alkaline phosphatases and CM-cellulase. There was a
significant effect of clay content on the concentrations of carbohydrates and activities of enzymes. The con-
centrations of carbohydrates increased when the clay contents of soils increased from 0 to 5 and 10%, respec-
tively, showing that the clay contents influence the capacity of soils for stabilization of carbohydrates. But
opposite trends were found in enzyme activities. The enzyme activities decreased significantly as the clay
contents of the artificial soils increased. The concentrations of carbohydrates and activities of enzymes were
significantly affected by exchangeable cations. In soils with 5 and 10% clay, the concentrations of carbohydrates
were maximum in Al-soils and minimum in Ca-soils, in contrast, the activities of enzymes were maximum in Ca-
soils and minimum in Al-soils. There was a significant effect of clay mineralogy on the concentrations of car-
bohydrates and activities of enzymes. The concentrations of carbohydrates were highest in soils with Wyoming
montmorillonite clay mineral and lowest in soils with Georgia kaolinite clay mineral. But the activities of en-
zymes were lower in soils with Wyoming montmorillonite clay mineral than soils with Georgia kaolinite and
Illinois illite clay minerals, indicating the influence of specific surface area (SSA) and cation exchange capacity
(CEC) of clay minerals on carbohydrate retention and enzyme activities. The results of this study indicate that
carbohydrates are stabilized in soils through the interaction with clay minerals and a small amount of clay (5%)
significantly increases carbohydrate retention and reduces enzyme activities in soils. Exchangeable cations exert
their influence on enzyme activities and hence carbohydrate dynamics by controlling the activities of enzymes
through modifying the physicochemical characteristics of soils.
1. Introduction
Better understanding of carbon cycling is necessary to decrease the
emission of greenhouse gases and global climate changes resulted from
it. Soil organic carbon (SOC) is a significant pool that plays important
role in global carbon cycling (Scharlemann et al., 2014; Wang and
Hsieh, 2002). The SOC consists of two main fractions; the labile and the
non-labile organic fractions. Carbohydrate constitutes an important
component of the labile fraction that represents about 5–25% of soil
organic matter (SOM) (Uzoho and Igbojionu, 2014). Its fate in soils
varies depending on the SOM dynamics and transformation processes,
especially decomposition, mineralization and immobilization (Paul
et al., 2002). Carbohydrate plays fundamental roles in the soil which
include; conservation of soil physical properties (Martins et al., 2012),
stabilization of soil aggregates (Yousefi et al., 2008) and sustenance of
soil microbial activities through the provision of readily available en-
ergy (Ros et al., 2003).
Besides being a source of energy for microorganisms, the soil car-
bohydrates play a significant role in the formation of stable aggregates
and metal ion complexes (Six et al., 2004). Soil carbohydrates consist of
plant- and microbial-derived polysaccharides that are an immediate by-
product of microbial utilization of organic carbon (OC). Microbial ex-
tracellular polysaccharides are regarded important in ecophysiological
processes from the viewpoints of resistance to desiccation and survival
https://doi.org/10.1016/j.clay.2018.07.031
Received 25 March 2018; Received in revised form 9 July 2018; Accepted 19 July 2018
⁎
Corresponding author.
E-mail address: rakhsh.fatemeh@znu.ac.ir (F. Rakhsh).
Applied Clay Science 163 (2018) 214–226
0169-1317/ © 2018 Elsevier B.V. All rights reserved.
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