Tree species identity alters decomposition of understory litter and associated microbial communities : a case study

Investigations on how tree species modify decomposition of understory litter have rarely been conducted, although potentially having impacts on soil carbon stocks and stability. The aim of our study was to disentangle the effects different tree species (alder, spruce, oak, and willow) exert on litter decomposition by comparing decomposition patterns and microbial measures (phospholipid fatty acids and microbial DNA) of both tree and understory (Calamagrostis epigejos) litter exposed at the respective tree species stands of a common garden experiment. An initially uniform mass loss of understory litter exposed at the stands suggests that inherent litter quality (assessed by C:N ratios and lignin content) was the major driver in early decomposition. However, in later stages of our experiment, decomposition of understory litter began to differ among the stands, suggesting a delayed tree species effect. Here, differences in microbial community composition caused by tree species identity (e.g., through varying N supply or phenolics leached from low-quality litter) were likely the major determinants affecting the decomposition of understory litter. However, in these advanced decomposition stages, tree species identity only partly altered microbial communities associated with understory litter. These results indicate that the development of microbial communities on understory litter (and its decay) is likely a combined result of inherent chemical composition and tree species identity.Peer reviewe

Unlocking Complex Soil Systems as Carbon Sinks: Multi-pool Management as the Key

Much research focuses on increasing carbon storage in mineral-associated organic matter (MAOM), in which carbon may persist for centuries to millennia. However, MAOM-targeted management is insufficient because the formation pathways of persistent soil organic matter are diverse and vary with environmental conditions. Effective management must also consider particulate organic matter (POM). In many soils, there is potential for enlarging POM pools, POM can persist over long time scales, and POM can be a direct precursor of MAOM. We present a framework for context-dependent management strategies that recognizes soils as complex systems in which environmental conditions constrain POM and MAOM formation

From fibrous plant residues to mineral-associated organic carbon – the fate of organic matter in Arctic permafrost soils

Permafrost-affected soils of the Arctic account for 70 % or 727 Pg of the soil organic carbon (C) stored in the northern circumpolar permafrost region and therefore play a major role in the global C cycle. Most studies on the budgeting of C storage and the quality of soil organic matter (OM; SOM) in the northern circumpolar region focus on bulk soils. Thus, although there is a plethora of assumptions regarding differences in terms of C turnover or stability, little knowledge is available on the mechanisms stabilizing organic C in Arctic soils besides impaired decomposition due to low temperatures. To gain such knowledge, we investigated soils from Samoylov Island in the Lena River delta with respect to the composition and distribution of organic C among differently stabilized SOM fractions. The soils were fractionated according to density and particle size to obtain differently stabilized SOM fractions differing in chemical composition and thus bioavailability. To better understand the chemical alterations from plant-derived organic particles in these soils rich in fibrous plant residues to mineral-associated SOM, we analyzed the elemental, isotopic and chemical composition of particulate OM (POM) and clay-sized mineral-associated OM (MAOM). We demonstrate that the SOM fractions that contribute with about 17 kg C m$^{-3}$ for more than 60 % of the C stock are highly bioavailable and that most of this labile C can be assumed to be prone to mineralization under warming conditions. Thus, the amount of relatively stable, small occluded POM and clay-sized MAOM that currently accounts with about 10 kg C m$^{-3}$ for about 40 % of the C stock will most probably be crucial for the quantity of C protected from mineralization in these Arctic soils in a warmer future. Using δ$^{15}$N as a proxy for nitrogen (N) balances indicated an important role of N inputs by biological N fixation, while gaseous N losses appeared less important. However, this could change, as with about 0.4 kg N m$^{-3}$ one third of the N is present in bioavailable SOM fractions, which could lead to increases in mineral N cycling and associated N losses under global warming. Our results highlight the vulnerability of SOM in Arctic permafrost-affected soils under rising temperatures, potentially leading to unparalleled greenhouse gas emissions from these soils

Abundance of lipids in differently sized aggregates depends on their chemical composition

Evidence for a vital role of soil mineral matrix interactions in lipid preservation is steadily increasing. However, it remains unclear whether solvent-extractable (‘free’) or hydrolyzable (‘bound’) lipids, including molecular proxies, e.g., for cutin and suberin, are similarly affected by different stabilization mechanisms in soil (i.e., aggregation or organo-mineral association). To provide insights into the effect of these stabilization mechanisms on lipid composition and preservation, we investigated free and bound lipids in particulate and mineral soil fractions, deriving from sand- and silt-/clay-sized aggregates from a forest subsoil. While free lipids accumulated in sand-sized aggregates, the more complex bound lipids accumulated in silt- and clay-sized aggregates, particularly in the respective mineral fractions < 6.3 µm (fine silt and clay). The presence of both, cutin and suberin markers indicated input of leaf- and root-derived organic matter to the subsoil. Yet, our cutin marker (9,10,ω-trihydroxyoctadecanoic acid) was not extracted from the mineral aggregate compartments < 6.3 µm, perhaps due to its chemical structure (i.e., cross-linking via several hydroxy groups, and thus higher ‘stability’, in macromolecular structures). Combined, these results suggest that the chemical composition of lipids (and likely also that of other soil organic matter compounds) governs interaction with their environment, such as accumulation in aggregates or association with mineral soil compartments, and thus indirectly influences their persistence in soil

Abundance of lipids in differently sized aggregates depends on their chemical composition

Evidence for a vital role of soil mineral matrix interactions in lipid preservation is steadily increasing. However, it remains unclear whether solvent-extractable (‘free’) or hydrolyzable (‘bound’) lipids, including molecular proxies, e.g., for cutin and suberin, are similarly affected by different stabilization mechanisms in soil (i.e., aggregation or organo-mineral association). To provide insights into the effect of these stabilization mechanisms on lipid composition and preservation, we investigated free and bound lipids in particulate and mineral soil fractions, deriving from sand- and silt-/clay-sized aggregates from a forest subsoil. While free lipids accumulated in sand-sized aggregates, the more complex bound lipids accumulated in silt- and clay-sized aggregates, particularly in the respective mineral fractions < 6.3 µm (fine silt and clay). The presence of both, cutin and suberin markers indicated input of leaf- and root-derived organic matter to the subsoil. Yet, our cutin marker (9,10,ω-trihydroxyoctadecanoic acid) was not extracted from the mineral aggregate compartments < 6.3 µm, perhaps due to its chemical structure (i.e., cross-linking via several hydroxy groups, and thus higher ‘stability’, in macromolecular structures). Combined, these results suggest that the chemical composition of lipids (and likely also that of other soil organic matter compounds) governs interaction with their environment, such as accumulation in aggregates or association with mineral soil compartments, and thus indirectly influences their persistence in soil

Preferential degradation of leaf- vs. root-derived organic carbon in earthworm-affected soil

Earthworms are integral parts of many ecosystems and may play a decisive role in determining whether soils function as carbon (C) sink or source. However, information on how earthworms affect the composition and stability of soil organic matter (SOM) is scarce. Particularly their effect on organic matter deriving from leaves and roots with distinct composition and, thus, susceptibility to decomposition and stabilization remains unclear. Here, we combine cutin- and suberin-derived lipids as specific markers for leaf- and root-derived SOM with their 13C composition and physical fractionations of soil. We show that earthworms overprint the protective role of organo-mineral associations and aggregates to favor the accumulation of root- relative to leaf-derived SOM. This gradual accumulation contributes to the often-observed dominance of root-derived organic matter in soil and emphasizes the need to consider molecular level effects of earthworms on SOM dynamics

Plant- or microbial-derived? A review on the molecular composition of stabilized soil organic matter

Soil organic matter (SOM) represents a major reservoir of stored carbon (C). However, uncertainties regarding the composition and origin of stabilized SOM hinder the implementation of sustainable management strategies. Here, we synthesize data on the contribution of plant- and microbial-derived compounds to stabilized SOM, i.e., aggregates and mineral-associated organic matter (MAOM), and review the role of environmental factors influencing this contribution. Extrapolating amino sugar concentrations in soil based on molecular stoichiometry, we find that microbial necromass accounts for ~50% (agroecosystems) or less (forest ecosystems) of the C stabilized within aggregates and MAOM across studies. This implies that plant biomolecules, including lipids, lignin, and sugars, might account for a substantial portion (≥50%) of the organic matter protected by minerals and aggregates. Indeed, plant-specific sugars and lipids can each account for as much as 10% of organic C within mineral soil fractions, and most reported quantities of plant-specific lipids and lignin in mineral soil fractions are likely underestimates due to irreversible sorption to minerals. A relatively balanced contribution of plant and microbial biomolecules to stabilized SOM in aggregates and MAOM is inconsistent with recent suggestions that stable SOM is comprised mostly of microbial compounds. Land use and soil type appear to profoundly affect the contribution of plant and microbial compounds to stabilized SOM. Consistent with studies of bulk soils, favorable conditions for microbial proliferation in grasslands or fertile Chernozems or Luvisols appear to increase the contribution of microbial compounds, while less favorable conditions for microbial proliferation in forest soils or Podzols/Alisols appear to favor the abundance of plant compounds in stabilized SOM. Combined with a tight link between substrate quality and the abundance of microbial compounds in stabilized SOM, and a potentially inverse relationship between substrate quality and the abundance of plant compounds, these results provide evidence that plant biomolecules might be preferentially stabilized by organo-mineral interactions in some ecosystems. Various areas warrant further research. For example, difficulties in distinguishing direct and indirect effects of temperature and precipitation on the composition of stabilized SOM may be overcome by long-term observational studies that include climate manipulations. Knowledge gaps in the contribution of plant and microbial compounds to stabilized SOM in soil layers below 30 cm depth may simply be closed by extending the sampling depth. Moreover, a refined focus on soil fauna, with potentially strong effects on microbial and plant compounds in stabilized SOM, will provide new insights into SOM dynamics. Future studies should quantify both microbial and plant biomolecules in mineral soil fractions to allow direct comparisons and overcome limitations in existing data. For example, because biomarker-based estimates of microbial-derived C can only indirectly estimate the maximum amount of plant-derived C, exhaustive studies of plant biomarker concentrations could be conducted, including estimates of plant-specific lipids, sugars, and lignin (and biomarkers released following mineral dissolution). Generally, more integrative studies, e.g., combining molecular and isotopic tracers of organic matter inputs with targeted sampling of mineral fractions, are required to improve knowledge of the formation and persistence of stabilized SOM

Tracing the sources and spatial distribution of organic carbon in subsoils using a multi-biomarker approach

Soil organic carbon (SOC) from aboveground and belowground sources has rarely been differentiated although it may drive SOC turnover and stabilization due to a presumed differing source dependent degradability. It is thus crucial to better identify the location of SOC from different sources for the parameterization of SOC models, especially in the less investigated subsoils. The aim of this study was to spatially assess contributions of organic carbon from aboveground and belowground parts of beech trees to subsoil organic carbon in a Dystric Cambisol. Different sources of SOC were distinguished by solvent-extractable and hydrolysable lipid biomarkers aided by C-14 analyses of soil compartments <63 mu m. We found no effect of the distance to the trees on the investigated parameters. Instead, a vertical zonation of the subsoil was detected. A high contribution of fresh leaf-and root-derived organic carbon to the upper subsoil (leaf-and root-affected zone) indicate that supposedly fast-cycling, leaf-derived SOC may still be of considerable importance below the A-horizon. In the deeper subsoil (root-affected zone), roots were an important source of fresh SOC. Simultaneously, strongly increasing apparent C-14 ages (3860 yrs BP) indicate considerable contribution of SOC that may be inherited from the Pleistocene parent material

Soil texture affects the coupling of litter decomposition and soil organic matter formation

Incomplete knowledge on the environmental factors linking litter decomposition and the formation of soil organic matter (SOM) hampers the sustainable management of soil as a carbon (C) sink. Here, we explored the effect of soil texture on the fate of C from decomposing litter (Indiangrass; Sorghastrum nutans (L.) Nash) and the concurrent formation of SOM in mineral soils of different textures (sand- and clay-rich) and forest floor material. We quantified the amount of litter C respired, C remaining in the litter, and litter C retained in the soil/forest floor in a 186-day incubation employing stable isotope analyses (C-13). We complemented our isotopic approach with the extraction of microbial biomarkers from the litter and soils/forest floor material and spectroscopic studies into the compositional changes of the incubated materials. We found that soil texture affected both the decomposition of litter and the retention of litter-derived C in the soil. The soil rich in clay provided conditions favorable for a more efficient microbial utilization of the litter material (high pH and high C use efficiency) as compared to the sand-rich soil and the forest floor. This resulted in lower amounts of litter C respired as CO2 (25.0%, vs. 55.6 and 56.1% in clay vs. sand and forest floor material, respectively) and higher amounts of litter C retained in the clay-rich soil (12.6% vs. 3.5 and 5.3% in clay vs. sand and forest floor material, respectively). High contents of silt- and clay-sized mineral particles in the clay-rich soil likely resulted in the ability to stabilize litter C in aggregates and organo-mineral associations, perhaps as microbial residues. This ability was low in the sand-rich soil and virtually absent in the forest floor, where the recalcitrance of the litter and native SOM was probably more relevant, and a larger portion of litter C may have been retained in the soil as relatively untransformed plant compounds. We emphasize that litter decomposition, the formation of SOM, and soil texture are tightly linked, such that any differences in soil texture alter litter decomposition and SOM formation patterns for the same litter.Peer reviewe