what does organic matter do to soil density
Introduction
Despite comprising a small proportion of the mass of agricultural soils, soil organic thing (SOM) is associated with improved soil structure (Feller and Beare, 1997; Six et al., 2000; Dexter et al., 2008; King et al., 2019). Multiple features of agricultural systems limit the extent by which direction tin can alter SOM levels, simply managing for fifty-fifty a modicum of increased SOM offers societal benefits, including climate change mitigation from the storage of carbon (C; Paustian et al., 2016; Minasny et al., 2017), a component of SOM, and reduced erosion (Barthès and Roose, 2002). Another, potential benefit of SOM is increasing crop yield (Pan et al., 2009; Oldfield et al., 2018), nonetheless, we take express ability to explain inconsistent effects of direction-induced increases in SOM on crops (Xin et al., 2016; Bradford et al., 2019; Wade et al., 2020) or to constrain potential crop benefits from SOM in future climates, as ingather stressors shift (IPCC, 2019). Progress on these fronts relies on a audio understanding of mechanisms through which SOM benefits crops in the first place.
Historically, improved ingather yield from SOM has primarily been attributed to SOM's office in resources supply, either of water or nitrogen (Gregorich et al., 1994; Arshad and Martin, 2002; Lal, 2020). A big effect of SOM on available h2o capacity (AWC) was established without consideration of limits of management-induced SOM (Hudson, 1994). A recent synthesis, however, concluded that management-induced increases in SOM (10 g C/kg soil) effected on average merely one.16 mm additional AWC in the top 10 cm of soil (Minasny and McBratney, 2018). This is a express contribution to ingather transpiration, which can exceed 450 mm (Kimball et al., 2019, maize). If a 1.sixteen mm increase in SOM-derived AWC is multiplied by rain events during crop maturation, signifying the number of times AWC is "used," and including 10–20 cm soil, the augmentation of AWC by SOM is larger, only an effect on ingather performance is likely context-dependent. While SOM is linked to AWC and management increases AWC in some situations, crop water supply does not announced well-justified as a universal machinery linking management-induced SOM increases to crop yield.
Larger SOM pools are linked to higher production rates of found-available nitrogen (N) via net North mineralization (Schimel, 1986; Berth et al., 2005), leading to the view that SOM benefits crops by increasing N supply. Withal, to determine that Northward supply from SOM limits crop growth, rates of plant Due north uptake should approach rates of N mineralization, indicating a potential for ingather Due north demand that outpaces soil N supply. Results of this comparison depend on method, with net Northward mineralization deceeding (Brye et al., 2003; Loecke et al., 2012) but gross North mineralization exceeding (Osterholz et al., 2016) crop N need. If gross Northward mineralization is indeed a improve indicator of plant-bachelor N than net North mineralization (Schimel and Bennett, 2004), or an underestimate given crop uptake of amino acids (Loma et al., 2011), then increasing SOM would merely be increasing an already sufficient N supply.
Here, we posit that SOM benefits crops non necessarily past increasing resource supply only by catalyzing crop resources capture (Figure i). Roots, the locus of food and h2o uptake, rely on contact with soil for nutrient (Wang et al., 2006), and water uptake (Javot and Maurel, 2002). Vigorous root evolution therefore multiplies the area of root-soil contact through which resource uptake occurs. We highlight below the consequences of inadequate aeration and compaction on roots and the ability of SOM-enhancing management practices to alleviate these stressors. While nosotros acknowledge that relationships betwixt SOM and soil construction are known (Karlen et al., 2001; Bünemann et al., 2018), we fence that their connections to related crop stressors—poor aeration and compaction—have received inadequate attending equally mechanisms that link SOM to crops. We also admit that compaction and inadequate aeration are often co-located, and nosotros separate them to talk over how they manifest differently in the soil environment and as root stressors.
Figure 1. Conceptual models representing (A) historically acknowledged and (B) newly proposed mechanisms linking management-induced increases in soil organic matter with crop yield. We recognize that the effect of SOM in mediating soil structure has been known for decades, but nosotros argue that exploring the response of crops to related stressors (inadequate aeration and compaction) has potential to explain effects of SOM on crop yield.
We refer to differences in SOM induced by agricultural practices, such as cover crops (Poeplau and Don, 2015), perennial forages (Male monarch and Blesh, 2018), manure application (Maillard and Angers, 2014), straw memory (Liu et al., 2014), or reduced tillage (in surface soil, Luo et al., 2010), except where noted. Although some of the mechanisms discussed may utilise to naturally occurring variability in SOM, we defer their discussion for brevity.
Aeration
Inadequate Aeration in Waterlogged and Non-saturated Soils
Equally a substrate for respiration, O2 and its transport potentially bear upon all functions of crop roots (Grable, 1966). To study the effects of Oii deficiency in the field, researchers commonly impose an extreme constriction of aeration with waterlogging, the saturation of soil pores with h2o (Hodgson and Chan, 1982; Bange et al., 2004). Our consideration of inadequate aeration focuses accordingly on waterlogging, and we do non review impacts of inadequate aeration in not-saturated soils on crops, equally these remain largely unexplored.
Waterlogging, Even if Transient, Tin can Indelibly Damage Roots
Waterlogging damages diverse crops worldwide (Velde and Van Der Tubiello, 2012; Shaw et al., 2013; Zhang et al., 2015; Li et al., 2019), and increases in waterlogging due to climate change are forecasted to exacerbate these damages (Rosenzweig et al., 2002). Waterlogging reduces soil aeration due to a 10iv fold slower rate of O2 diffusion through water than through air (Grable, 1966). Inside hours, soil Oii tin can be depleted as root or soil organisms' respiration demands exceed atmospheric O2 supply, with rate and extent of Oii depletion depending on depth (Malik et al., 2001) and temperature (Trought and Drew, 1982). Over slightly longer fourth dimension frames, waterlogged soils also accumulate byproducts of root or microbial metabolism, east.g., CO2, Fe2+, and Mntwo+, potential plant toxins (Shabala, 2011). Crop responses to waterlogging depend on crop species and cultivar (Huang, 1997; Boru, 2003; Ploschuk et al., 2018), every bit well equally timing (Rhine et al., 2010; de San Celedonio et al., 2014; Ren et al., 2014) and duration (Malik et al., 2002; Rhine et al., 2010; Kuai et al., 2014; Ren et al., 2014; Arduini et al., 2019) of waterlogging, but a well-supported view holds that ingather yield reductions from waterlogging are largely owing to impaired function and inadequate recovery of roots (Malik et al., 2002; Herzog et al., 2016; Arduini et al., 2019).
In crops that are susceptible to waterlogging, stress response can be considered both during and subsequently release from waterlogging. During waterlogging, O2 deficiency induces an energy crisis in the root due to inefficient production of ATP (Gibbs and Greenway, 2003), which curtails energy-dependent nutrient uptake (Trought and Drew, 1980; Morard et al., 2000; Colmer and Greenway, 2011) and root growth (Palta et al., 2010; Arduini et al., 2019). After release from waterlogging, root growth of certain root forms, i.e., seminal roots, can be permanently inhibited (Malik et al., 2002; Palta et al., 2010; Colmer and Greenway, 2011), attributable to cell death in upmost meristems (Trought and Drew, 1980; Malik et al., 2002), beyond the attain of establish-transported O2 (Colmer and Greenway, 2011). Crops must then rely on energetically-expensive production of new roots, i.eastward., adventitious roots (Palta et al., 2010; Steffens and Rasmussen, 2016), and/or increase nutrient uptake per unit root (Arduini et al., 2019). These adaptations practise not necessarily let crops to escape a waterlogging yield penalty, equally reductions in root biomass (Grassini et al., 2007; de San Celedonio et al., 2017; Ploschuk et al., 2018) or root length density (Hayashi et al., 2013) are oft linked to reductions in shoot biomass or yield.
Doubtfulness remains about the threshold duration at which waterlogging damages crops. In some crops and growth stages, waterlogging equally short equally 3 days reduced yield (Malik et al., 2002; Ren et al., 2014), simply the shortest waterlogging we constitute in field studies was 2 days (Rhine et al., 2010). Due to the risk of crop damage from fifty-fifty transient waterlogging, there is interest in management practices that reduce waterlogging hazard (Manik et al., 2019) and better root aeration across the soil h2o spectrum (Rabot et al., 2018).
Soil Organic Matter: Means to Improve Root Aeration
Reducing Duration of Waterlogging
Management practices that promote SOM reduce risk and duration of waterlogging by increasing rate of water infiltration (Boyle et al., 1989; Adekalu et al., 2007; Abid and Lal, 2009; Blanco-Canqui et al., 2011), which increases the time soil can receive pelting before ponding occurs (McGarry et al., 2000) and reduces time required to drain from saturation to field chapters (Wuest et al., 2005). Among the many measurable soil h2o variables, infiltration is the nearly usually assessed, and we notation the need to ameliorate establish relationships betwixt infiltration, time to ponding, and drainage. Information technology is also important to note, as reviewed by Blanco-Canqui and Ruis (2018) for no-till, that direction practices that promote SOM can take a neutral effect on water infiltration in some cases, despite positive effects in majority of cases.
Accelerated infiltration associated with SOM is attributable to several soil features. The redistribution of soil mass to larger aggregate size classes associated with SOM (King et al., 2019) helps to explain an increase in total (Pikul and Zuzel, 1994; Yang et al., 2011; Blanco-Canqui and Benjamin, 2013) or macro- porosity (>0.3–0.4 mm, Deurer et al., 2009; Yagüe et al., 2016), although this effect is non detectable in all cases (Ruiz-Colmenero et al., 2013). SOM also stabilizes aggregates (Chenu et al., 2000; Annabi et al., 2011), minimizing their dissolution into smaller, and pore-bottleneck size fractions that seal the soil surface against water infiltration (Bissonnais and Arrouays, 1997; Lado et al., 2004). The well-nigh dramatic effects of SOM on infiltration tin probable be traced to earthworms and/or termites and their cosmos of wide, continuous, vertically-oriented pores through which water flows preferentially (McGarry et al., 2000; Guo and Lin, 2018). More abundant – or more active (Pérès et al., 2010) – soil fauna may exist due in part to reduced disturbance associated with some SOM-promoting practices, e.g., no-till. Notwithstanding, close relationships between SOM and earthworm abundance without the confounding event of disturbance (Fonte et al., 2009; Guo et al., 2016) also indicate a part for SOM equally faunal substrate supply.
Few studies take attempted to link SOM-induced reductions in waterlogging with crop yield. Gómez-paccard et al. (2015), still, notice crop yields increased from reduced surface soil waterlogging associated with no-till. The power of SOM-mediated reductions in waterlogging to do good crops are most likely when (1) ingather is sensitive to waterlogging and (2) rainfall intensity can be mediated by SOM on a timescale relevant to waterlogging stress (neither drizzle nor deluge); and (3) soil is otherwise poorly-drained (Rhine et al., 2010).
Promoting Aeration in Non-saturated Soils
If crops feel inadequate aeration in non-saturated soils, it is reasonable to look that SOM would better gas diffusivity given its effects on related parameters of soil structure (Neira et al., 2015, and below). Few studies investigate the isolated effect of SOM on gas diffusivity, however, Colombi et al. (2019) discover a positive relationship between SOM and gas diffusivity at field capacity across a soil texture slope. Future piece of work should examine net effects of SOM on O2 diffusivity and consumption in soils.
Compaction
Soil Compaction Constrains Root Evolution
Soil compaction reduces crop yields (Coelho et al., 2000; Ishaq et al., 2001a; Bayhan et al., 2002; Czyz, 2004; Whalley et al., 2008) and is quantified via either bulk density or mechanical impedance (MI; Ehlers et al., 1987; Bengough et al., 2011). MI estimates the force encountered by the elongation of a living root, and is consequential for crops because greater MI inflates the photosynthate required for root elongation (Herrmann and Colombi, 2019). Although MI measurements ignore biopores used preferentially by roots (Stirzaker et al., 1996; White and Kirkegaard, 2010), MI is more descriptive than bulk density because it is sensitive to soil water. Drying soils present increasing MI (Vaz et al., 2011), and to isolate effects of water stress from compaction stress per se on crop evolution, researchers apply experimental compaction.
Compaction studies betoken that reduced crop yield from compaction is due in large part to constraints on root evolution (Ishaq et al., 2001b; Czyz, 2004; Colombi and Keller, 2019). A root restricted past soil compaction is generally thicker than a root in non-compacted soil (Nadian et al., 1997), likely due to greater axial strength needed to overcome compaction (Bengough, 2012). Reductions in total number of roots, charge per unit of root elongation, full root length, or root biomass are also reported (Panayiotopoulos et al., 1994; Chan et al., 2006; Lipiec et al., 2012). Root length is generally more reduced than root dry mass (Panayiotopoulos et al., 1994), indicating the aggregating of belowground photosynthate without commensurate expansion of soil-contacting surface area available for nutrient and water uptake. Compaction is sometimes characterized by a hardpan around twenty cm depth, which leads to restricted root access to subsoil and concentrated root development in the topsoil (Czyz, 2004). This pattern of root development prevents crop access of deep soil water most implicated in ingather drought resistance (Uga et al., 2013; Lynch, 2018).
A single threshold MI for ingather sensitivity is unlikely to serve universally, and not simply because cultivars (Houlbrooke et al., 1997) and crops (Rosolem et al., 2002) differ in MI tolerance. Threshold MI values also likely depend on definition by energy required to extend roots (Herrmann and Colombi, 2019) or past crop yield penalty. The MI required to reduce root growth efficiency is likely less than required to touch yields, and yield penalisation due to restricted roots can be counteracted somewhat by fertilization (Robertson et al., 2009). Whatever the threshold, the detrimental effects of compaction on crops has generated attention toward means to reduce it.
Soil Organic Thing Reduces Compaction and Is Associated With Root Channels
Reduced Compaction: More than Water Transpired Before Mechanical Impedance Limits Growth
Although SOM is often promoted for its ability to alleviate soil compaction and associated increases in MI (Hamza and Anderson, 2005), the generation of data confirming a negative SOM-MI human relationship has been hampered by the convention of measuring MI in soils nigh field capacity (Duiker, 2002). Even in large datasets, no relationship between SOM and MI in soil nearly field capacity is found (Fine et al., 2017), probable because very wet soils (∼1–ten kPa) offer minimal MI regardless of SOM. Information technology is as MI increases in drying soils (Vaz et al., 2011; Filho et al., 2014) that an effect of SOM becomes credible (Stock and Downes, 2008; Gao et al., 2012).
Nosotros highlight two studies showing the upshot of SOM in reducing MI as soils dry out. Stock and Downes (2008) and Gao et al. (2012) investigated soils differing just in SOM concentrations. With few exceptions, MI increased as soils approached permanent wilting signal, but the increment in MI was not as much in higher SOM soils. In other words, SOM allows soil to become drier before reaching a potentially root-constraining MI. For instance, Stock and Downes (2008) find an MI of 1.5 MPa is reached in the 1% OM soil at ∼-100 kPa, whereas the iii% OM soil reaches the same MI at about ∼-200 kPa. For these soils, the difference in volumetric h2o content between -100 and -200 kPa is ∼0.02 m–3 H2O m–three soil, or ∼5 mm of water when considered over the peak 25 cm. While Stock and Downes (2008) added organic amendments to glacial till to create fixed SOM percentages, their results resemble those of Gao et al. (2012), who compared fallow to grassland soils. The extent to which direction-induced SOM benefits crops via reductions in compaction likely vary with context, particularly those relevant to MI thresholds (run into section "Soil compaction constrains root development").
Root Channels to Subsoil Water
Although not connected to the physical or biological backdrop of SOM, management practices that promote SOM may likewise alleviate the furnishings of compaction by facilitating crop root access to the subsoil. Deep-rooted cover crops or perennial crops create root channels to subsoil (McCallum et al., 2004), which are used past subsequent greenbacks crops (Rasse and Smucker, 1998; Williams and Weil, 2004). Crops are most likely to benefit from these root channels if subsoil is compacted and if crops experience sufficient water stress for subsoil h2o stores to exist relevant.
Discussion and Outlook
In the historical conceptual model linking crop performance to SOM, SOM benefits crops primarily by supplying nitrogen and h2o. Hither we propose SOM equally a mediator of resource uptake via root growth. We note caveats to the proposed framework. The extent to which SOM affects food and water supply however merits research, and non all mechanisms discussed are contingent on increased SOM. We focus on MI to characterize compaction, simply SOM may alter soil construction in means relevant to root growth that are non captured by MI. As a simple diagram, Figure 1 does non describe that yield penalty in low SOM soils may be due to toll of constructing new or thicker roots. However, considering SOM as catalyzing resource uptake via root development tin help explain recent reports of SOM furnishings on crops. Wade et al. (2020) study maize yield increases from direction-induced SOM across a range of N fertilizer levels, consistent with the concept that root N uptake—as well as soil Northward supply—can limit crop yield. Recognizing the importance of aeration for roots may too explain a parabolic crop response to a SOM gradient in a pot study (Oldfield et al., 2020), in which synthesized mixtures of minerals and organic horizons college in SOM may have supported college O2 consumption by microbes without improved gas diffusivity expected in natural loftier-SOM soils (Colombi et al., 2019).
We propose the exploration of SOM as a catalyst for resource capture focuses on:
• Characterizing the context-mediated issue of SOM on aeration and compaction by describing the effect of SOM gradients on (a) aeration and hazard and duration of waterlogging and (b) MI across a range of soil moisture contents.
• Investigating ingather response to SOM every bit a role of aeration and compaction by identifying the thresholds of hypoxia affecting roots and yields. To chronicle SOM-mediated MI to root development, describing soil moisture status during crop maturation will exist crucial.
• Simulating future crop response to SOM; currently, Basche et al. (2016) and Jarecki et al. (2018) are two of few examples to model the potential for management-induced SOM to stabilize crop yields under time to come climates.
We hope the lens proposed will assistance illuminate the effect of SOM on crops.
Author Contributions
AK conceptualized and drafted the manuscript. GA, AG, and CW-R provided input to arrive at the final version.
Funding
This work was funded past a Natural Sciences and Technology Enquiry Quango of Canada Discovery Grant to CW-R.
Conflict of Interest
The authors declare that the research was conducted in the absence of any commercial or fiscal relationships that could exist construed as a potential disharmonize of involvement.
Acknowledgments
We thank Peter Pellitier for feedback on an early on version of the manuscript and Rebecca Johnson for comments that improved the figure. Two reviewers identified careful distinctions in the literature review.
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