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208 records · Page 12Linked to original sources

Correlating sea lamprey density with environmental DNA detections in the lab

Invasive sea lamprey (Petromyzon marinus Linnaeus, 1758) are currently managed by the Great Lakes Fishery Commission in an effort to reduce pest populations below levels that cause ecological damage. One technique to improve stream population assessments could be molecular surveillance in the form of environmental DNA (eDNA) monitoring. We developed and validated four probe-based quantitative polymerase chain reaction (qPCR) assays, then used two probes (cytb, nd1) to determine whether eDNA concentration was correlated with adult and larval sea lamprey density in the lab. We found a strong positive correlation between adult sea lamprey densities of 2, 20, and 200 individuals/2000L and eDNA concentrations in tanks using both assays (cytb, nd1). For larval laboratory tank density trials, eDNA concentrations were generally near our limit of quantification and there was no significant difference in copy numbers detected between larval sea lamprey densities of 1, 5, and 25 individuals/28L. Therefore, we examined detection probability rather than concentration with laboratory tank densities. We observed a trend of increasing detection probabilities with increased larval sea lamprey density that approached significance suggesting that when DNA copy numbers are low, detection rates may be more informative in predicting varying densities of larval sea lamprey. The ability to assess sea lamprey densities from a water sample could be a powerful tool to improve traditional assessment and stream ranking techniques. Further refinement of this method in the field may make eDNA surveillance of sea lamprey a reliable part of stream assessments. Rapid eDNA analysis from many streams may help focus traditional assessment efforts, thereby improving the efficiency of invasive sea lamprey control efforts.

Management of Biological Invasions

Juvenile African clawed frogs (Xenopus laevis) express growth, metamorphosis, mortality, gene expression, and metabolic changes when exposed to thiamethoxam and clothianidin

Neonicotinoids (NEO) represent the main class of insecticides currently in use, with thiamethoxam (THX) and clothianidin (CLO) primarily applied agriculturally. With few comprehensive studies having been performed with non-target amphibians, the aim was to investigate potential biomarker responses along an adverse outcome pathway of NEO exposure, whereby data were collected on multiple biological hierarchies. Juvenile African clawed frogs, Xenopus laevis , were exposed to commercial formulations of THX and CLO at high (100 ppm) and low (20 ppm) concentrations of the active ingredient. Mortality, growth, development, liver metabolic enzyme activity, and gene expression endpoints were quantified. Tadpoles ( n > 1000) from NF 47 through tail resorption stage (NF 66) were exposed to NEO or to NEO-free media treatments. Liver cell reductase activity and cytotoxicity were quantified by flow cytometry. Compared to control reference gene expressions, levels of expression for NEO receptor subunits, cell structure, function, and decontamination processes were measured by RT-qPCR by using liver and brain. Mortality in THX high was 21.5% compared to the control (9.1%); the metabolic conversion of THX to CLO may explain these results. The NF 57 control tadpoles were heavier, longer, and more developed than the others. The progression of development from NF 57–66 was reduced by THX low, and weight gain was impaired. Liver reductases were highest in the control (84.1%), with low NEO exhibiting the greatest reductions; the greatest cytotoxicity was seen with THX high. More transcriptional activity was noted in brains than in livers. Results affirm the utility of a study approach that considers multiple complexities in ecotoxicological studies with non-target amphibians, underscoring the need for simultaneously considering NEO concentration-response relationships with both whole-organism and biomarker endpoints.

International Journal of Molecular Sciences

The role of neutral and adaptive genomic variation in population diversification and speciation in two ground squirrel species of conservation concern

Understanding the neutral (demographic) and adaptive processes leading to the differentiation of species and populations is a critical component of evolutionary and conservation biology. In this context, recently diverged taxa represent a unique opportunity to study the process of genetic differentiation. Northern and southern Idaho ground squirrels ( Urocitellus brunneus —NIDGS, and U . endemicus —SIDGS, respectively) are a recently diverged pair of sister species that have undergone dramatic declines in the last 50 years and are currently found in metapopulations across restricted spatial areas with distinct environmental pressures. Here we genotyped single-nucleotide polymorphisms (SNPs) from buccal swabs with restriction site-associated DNA sequencing (RADseq). With these data we evaluated neutral genetic structure at both the inter- and intraspecific level, and identified putatively adaptive SNPs using population structure outlier detection and genotype–environment association (GEA) analyses. At the interspecific level, we detected a clear separation between NIDGS and SIDGS, and evidence for adaptive differentiation putatively linked to torpor patterns. At the intraspecific level, we found evidence of both neutral and adaptive differentiation. For NIDGS, elevation appears to be the main driver of adaptive differentiation, while neutral variation patterns match and expand information on the low connectivity between some populations identified in previous studies using microsatellite markers. For SIDGS, neutral substructure generally reflected natural geographical barriers, while adaptive variation reflected differences in land cover and temperature, as well as elevation. These results clearly highlight the roles of neutral and adaptive processes for understanding the complexity of the processes leading to species and population differentiation, which can have important conservation implications in susceptible and threatened species.

Idaho

Nuclear eDNA estimates population allele frequencies and abundance in experimental mesocosms

Advances in environmental DNA (eDNA) methodologies have led to improvements in the ability to detect species and communities in aquatic environments, yet the majority of studies emphasize biological diversity at the species level by targeting variable sites within the mitochondrial genome. Here, we demonstrate that eDNA approaches also have the capacity to detect intraspecific diversity in the nuclear genome, allowing for assessments of population-level genetic diversity and estimates of the number of genetic contributors in a sample. Using a panel of microsatellite loci, we evaluated intraspecific genetic diversity in the round goby (Neogobius melanostomus) using eDNA samples from experimental mesocosms. First, we tested the similarity between eDNA and individual tissue-based estimates of allele frequencies. Subsequently, we used a likelihood-based DNA mixture framework to estimate the number of unique genetic contributors in mesocosm eDNA samples and in simulated mixtures of alleles. Allele frequencies from eDNA accurately reflected allele frequencies from genotyped round goby tissue samples, indicating nuclear markers can be reliably amplified from water samples under controlled conditions. DNA mixture analyses were able to estimate the number of genetic contributors from eDNA samples and simulated mixtures of DNA from up to 58 individuals, with the degree of positive or negative bias dependent on the filtering scheme of low-frequency alleles. This study is the first to document the application of eDNA and multiple amplicon-based methods to obtain intraspecific nuclear genetic information and estimate the absolute abundance of a species in mesocosms. With proper validation, this approach has the potential to advance non-invasive survey methods to characterize populations and broadens the application of eDNA methodologies to inform population-level management objectives.

New York

Assessing grass carp (Ctenopharyngodon idella) occupancy and detection probability within Lake Erie from environmental DNA

Grass carp ( Ctenopharyngodon idella ), an invasive cyprinid within the Laurentian Great Lakes, is naturally reproducing in several Lake Erie tributaries, which has raised concerns of the species’ spread throughout Lake Erie and the other Great Lakes. Knowledge of the recent invasion extent outside of the western basin of Lake Erie, particularly in eastern tributaries and nearshore waters, is limited. Understanding the invasion extent would improve the efficacy of ongoing coordinated multi-agency control efforts. Molecular tools, such as environmental DNA (eDNA), have shown promise for early detection of aquatic invasive species. In this study, water samples (N = 476) were collected for grass carp eDNA monthly between May and November in 2018 and 2019, at three sites in the Michigan waters of Lake Erie and the Detroit River. We fit Bayesian multi-scale occupancy models to determine differences in eDNA capture and detection probability among grass carp qPCR assays, sampling sites, and across time. To determine whether grass carp were physically present, and to validate eDNA samples, we quantified recent grass carp presence in sampled areas using an existing acoustic telemetry and field sampling framework. Our results indicate that grass carp eDNA capture probability differed among sites, but there was no difference among months. Positive grass carp eDNA detections were observed across multiple months at each site, with 69% of site-specific sampling events testing positive for grass carp eDNA on at least one assay and replicate. The majority (65%) of weeks where positive eDNA sampling detections occurred also concurrently had one or more grass carp detected via acoustic telemetry 1–6 days prior. Our results highlight the potential utility of using eDNA to monitor the invasion extent of grass carp within the nearshore waters of Lake Erie. However, further evaluation of the factors that influence grass carp eDNA characteristics among sites within Lake Erie are needed to determine its efficacy for surveillance protocols by natural resource management agencies.

Lake Erie

A Cftr-independent, Ano1-rich seawater-adaptive ionocyte in sea lamprey gills

All ionoregulating marine fishes examined to date utilize seawater-type ionocytes expressing the apical Cl- channel, cystic fibrosis transmembrane conductance regulator (Cftr) to secrete Cl − . We performed transcriptomic, molecular, and functional studies to identify Cl − transporters in the seawater-type ionocytes of sea lamprey ( Petromyzon marinus ). Gill cftr expression was minimal or undetectable in larvae and post-metamorphic juveniles. We identified other Cl − transporters highly expressed in the gills and/or upregulated following metamorphosis and further investigated two candidates that stood out in our analysis, a Ca 2+ -activated Cl − channel, anoctamin 1 ( ano1 ), and the Clc chloride channel family member 2 ( clcn2 ). Of these, ano1 was expressed 10-100 times more than clcn2 in the gills; moreover, ano1 was upregulated during seawater acclimation, while clcn2 was not. Using an antibody raised against sea lamprey Ano1, we did not detect Ano1 in the gills of larvae, found elevated levels in juveniles and observed a 4-fold increase in juveniles after seawater acclimation. Ano1 was localized to seawater-type branchial ionocytes but, surprisingly, was localized to the basolateral membrane. In vivo pharmacological inhibition experiments demonstrated that a DIDS-sensitive mechanism was critical to the maintenance of osmoregulatory homeostasis in seawater- but not freshwater-acclimated sea lamprey. Taken together, our results provide evidence of a Cftr-independent mechanism for branchial Cl − secretion in sea lamprey that leverages Ano1-expressing ionocytes. Once further characterized, the Cftr-independent, Ano1-rich ionocytes of sea lamprey could reveal novel strategies for branchial Cl − secretion, whether by Ano1 or some other Cl − transporter, not previously known in ionoregulating marine organisms.

Journal of Experimental Biology

Chromosome painting in the manatee supports Afrotheria and Paenungulata

Background Sirenia (manatees, dugongs and Stellar's sea cow) have no evolutionary relationship with other marine mammals, despite similarities in adaptations and body shape. Recent phylogenomic results place Sirenia in Afrotheria and with elephants and rock hyraxes in Paenungulata. Sirenia and Hyracoidea are the two afrotherian orders as yet unstudied by comparative molecular cytogenetics. Here we report on the chromosome painting of the Florida manatee. Results The human autosomal and X chromosome paints delimited a total of 44 homologous segments in the manatee genome. The synteny of nine of the 22 human autosomal chromosomes (4, 5, 6, 9, 11, 14, 17, 18 and 20) and the X chromosome were found intact in the manatee. The syntenies of other human chromosomes were disrupted in the manatee genome into two to five segments. The hybridization pattern revealed that 20 (15 unique) associations of human chromosome segments are found in the manatee genome: 1/15, 1/19, 2/3 (twice), 3/7 (twice), 3/13, 3/21, 5/21, 7/16, 8/22, 10/12 (twice), 11/20, 12/22 (three times), 14/15, 16/19 and 18/19. Conclusion There are five derived chromosome traits that strongly link elephants with manatees in Tethytheria and give implicit support to Paenungulata: the associations 2/3, 3/13, 8/22, 18/19 and the loss of the ancestral eutherian 4/8 association. It would be useful to test these conclusions with chromosome painting in hyraxes. The manatee chromosome painting data confirm that the associations 1/19 and 5/21 phylogenetically link afrotherian species and show that Afrotheria is a natural clade. The association 10/12/22 is also ubiquitous in Afrotheria (clade I), present in Laurasiatheria (clade IV), only partially present in Xenarthra (10/12, clade II) and absent in Euarchontoglires (clade III). If Afrotheria is basal to eutherians, this association could be part of the ancestral eutherian karyotype. If afrotherians are not at the root of the eutherian tree, then the 10/12/22 association could be one of a suite of derived associations linking afrotherian taxa.

BMC Evolutionary Biology

Prevention, early detection and containment of invasive, nonnative plants in the Hawaiian Islands: current efforts and needs

Introduction: Invasive, non-native plants (or environmental weeds) have long been recognized as a major threat to the native biodiversity of oceanic islands (Cronk & Fuller, 1995; Denslow, 2003). Globally, several hundred non-native plant species have been reported to have major impacts on natural areas on oceanic islands (Kueffer et al ., 2009). In Hawaii, at least some 50 non-native plant species reach dominance in natural areas (Kueffer et al ., 2009) and many of them are known to impact ecosystem processes or biodiversity. One example is the invasive Australian tree fern ( Cyathea cooperi ), which has been shown to be very efficient at utilizing soil nitrogen and can grow six times as rapidly in height, maintain four times more fronds, and produce significantly more fertile fronds per month than the native Hawaiian endemic tree ferns, Cibotium spp. (Durand & Goldstein, 2001a, b). Additionally, while native tree ferns provide an ideal substrate for epiphytic growth of many understory ferns and flowering plants, the Australian tree fern has the effect of impoverishing the understory and failing to support an abundance of native epiphytes (Medeiros & Loope, 1993). Other notorious examples of invasive plant species problematic for biodiversity and ecosystem processes in Hawaii include miconia ( Miconia calvescens ), strawberry guava ( Psidium cattleianum ), albizia ( Falcataria moluccana ), firetree ( Morella faya ), clidemia ( Clidemia hirta ), kahili ginger ( Hedychium gardnerianum ), and fountain grass ( Pennisetum setaceum ), to name just a few. Fireweed ( Senecio madagascariensis ) is a recent example of a seriously problematic invasive species for Hawaii’s agriculture and is damaging certain high-elevations native ecosystems as well. The threat of invasive plants has long been recognized in Hawaii and is well documented (e.g. Cox, 1999; Loope & Kraus, 2009 in press; Loope et al ., 2004; Mooney & Drake, 1986; Stone & Scott, 1985; Stone et al. , 1992). In many respects, Hawaii may be near the forefront among national and international efforts to address the burgeoning threat of invasive plants, perhaps especially in the field of outreach and education (Holt, 1996; Van Driesche & Van Driesche, 2000). However, given the scale of the problem many challenges still need to be addressed and gaps in the existing management system need to be identified. In particular, it appears that new non-native plant species are still introduced to the Hawaiian Islands at a high rate with little or no regard for their potential invasiveness. In fact, a Pacific-wide and a global survey of non-native plants on oceanic islands have both shown that on Hawaii among all archipelagos by far the highest number of problematic invasive species known from other areas in the world is already present (Denslow et al . 2009, Kueffer et al . 2009). Hawaii lacks an effective mechanism for tracking what species are present or incoming. For instance, early detection nursery surveys conducted on Maui in 2008 found over 300 species of cultivated vascular plants that have not previously been recorded in Hawaii (Starr et al. , in prep.). In spite of an innovative Hawaii Biological Survey (e.g. Eldredge & Evenhuis, 2003), there is no mechanism for recording presence of a species until it becomes naturalized. Some of these new introductions may quickly become serious pests. Fireweed, first recorded in Hawaii on the Big Island in the early 1980s, is now considered one of the Kueffer & Loope 2009 5/48 worst weeds of pastures and is also invading natural areas from near sea level to above 10,000 feet. Although the cultivated and as yet non-invasive Cortaderia selloana has been present in Hawaii for 50 years or more, the morphologically similar Cortaderia jubata was simultaneously found to be present on Maui and invading on a large scale in 1989. It played an important role in inspiring the establishment of the Maui Invasive Species Committee (MISC) in 1997, and MISC now spends roughly $200,000 per year removing and containing C. jubata to keep it from becoming widespread in high elevation conservation lands of East and West Maui. The existence of many similar examples shows that to date regulatory action to prevent new invasive plant species from establishing and spreading in Hawaii has not yet been as successful as it needs to be. In particular, because some problematic invasive species known from other areas in the world (Kueffer et al ., 2009; Weber, 2003) have not yet been recorded from Hawaii, preventive measures against the introduction and spread of such likely invasive species is therefore an urgent need for Hawaii. Indeed, regulation of importation and early detection and eradication of introduced species before they become abundant and widespread are widely considered the most cost-efficient and often only effective measures against the threat of new invasive species (Kueffer & Hirsch Hadorn, 2008; Wittenberg & Cock, 2001). Timing seems favorable for Hawaii to achieve effective protection against the threat of new invasive species through prevention, early detection, and eradication/containment. Through the establishment and evolution of Invasive Species Committees (ISCs) on each major Hawaiian island, the institutional capacity has been built up for prevention, early detection, containment, and outreach at an island scale. Weed risk assessment (Daehler et al ., 2004) and early detection methodologies (Starr et al. , in review-a, b) have been developed and tested specifically for Hawaii. Containment strategies have been successful (e.g., Special Ecological Areas in Hawaii Volcanoes National Park), and so have eradications of particular species on an island scale (e.g. mullein ( Verbascum thapsus ) and other species on Maui, fireweed ( Senecio madagascariensis ) on Kauai). These successful management strategies may be further strengthened through recently developed novel approaches in research (e.g. remote sensing, species distribution modelling, and molecular genetics tools). Another major recent achievement is the gained support of the plant industry for preventive measures against invasive species (see p. 13ff). Last but not least, regulatory action is also moving forward. Passage of House Bill 2517 by the 2008 Hawaii House and Senate and prompt signing of the bill into law by the Governor provides hope that action to ban the sale of a meaningful suite of restricted weeds can quickly proceed through the rulemaking phase into the implementation phase. This report documents these achievements and experiences and provides a range of perspectives on how to further develop prevention, early detection and containment of invasive species in Hawaii. The report is based on a symposium and workshop held at the 2008 Hawaii Conservation Conference in Honolulu on 31 July 2008.

Hawaii

Abundance estimation and conservation biology

Abundance is the state variable of interest in most population–level ecological research and in most programs involving management and conservation of animal populations. Abundance is the single parameter of interest in capture–recapture models for closed populations (e.g., Darroch, 1958; Otis et al., 1978; Chao, 2001). The initial capture–recapture models developed for partially (Darroch, 1959) and completely (Jolly, 1965; Seber, 1965) open populations represented efforts to relax the restrictive assumption of population closure for the purpose of estimating abundance. Subsequent emphases in capture–recapture work were on survival rate estimation in the 1970’s and 1980’s (e.g., Burnham et al., 1987; Lebreton et al.,1992), and on movement estimation in the 1990’s (Brownie et al., 1993; Schwarz et al., 1993). However, from the mid–1990’s until the present time, capture–recapture investigators have expressed a renewed interest in abundance and related parameters (Pradel, 1996; Schwarz & Arnason, 1996; Schwarz, 2001). The focus of this session was abundance, and presentations covered topics ranging from estimation of abundance and rate of change in abundance, to inferences about the demographic processes underlying changes in abundance, to occupancy as a surrogate of abundance. The plenary paper by Link & Barker (2004) is provocative and very interesting, and it contains a number of important messages and suggestions. Link & Barker (2004) emphasize that the increasing complexity of capture–recapture models has resulted in large numbers of parameters and that a challenge to ecologists is to extract ecological signals from this complexity. They offer hierarchical models as a natural approach to inference in which traditional parameters are viewed as realizations of stochastic processes. These processes are governed by hyperparameters, and the inferential approach focuses on these hyperparameters. Link & Barker (2004) also suggest that our attention should be focused on relationships between demographic processes such as survival and recruitment, the two quantities responsible for changes in abundance, rather than simply on the magnitudes of these quantities. They describe a type of Jolly–Seber capture–recapture model that permits inference about the underlying relationship between per capita recruitment rates and survival rates (Link & Barker, this volume). Implementation used Bayesian Markov Chain Monte Carlo methods and appeared to work well, yielding inferences about the relationship between recruitment and survival that were robust to selection of prior distribution. We believe that readers will find their arguments compelling, and we expect to see increased use of hierarchical modeling approaches in capture–recapture and related fields. Otto (presentation without paper) also recommended use of hierarchical models in analysis of multiple data sources dealing with population dynamics of North American mallards. He integrated survival inferences from ringing data, abundance information from aerial survey data, and recruitment information based on age ratios from a harvest survey. He used a Leslie matrix population projection model as an integrating framework and obtained estimates of breeding population size using all data.Otto’s approach also permitted inference about biases in estimated quantities. As with the work of Link & Barker (2004), we find Otto’s recommendation to use hierarchical models to integrate data from multiple sources to be very compelling. Alisauskas et al. (2004) report results of an analysis of capture–recapture data for a askatchewan population of white–winged scoters. They used the approach of Pradel (1996) to estimate population growth rate (See the PDF) directly. Estimates for 1975–1985 were quite low, but estimates for the recent period, 2000–2003,increased to values > 1. Parameter estimates for seniority, survival and per capita recruitment (Pradel, 1996) led to the inference that increased recruitment was largely responsible for the improvements in population status and growth. However, various data sources also indicated that this increase in recruitment was likely a result of increased immigration rather than improved reproduction on the area. This latter inference is important from a conservation perspective in indicating the importance of birds in other locations to growth and health of the study population. Lukacs and Burnham presented material to be published elsewhere that dealt with the use of genetic markers in capture–recapture studies. The data sources for such studies are samples of hair or feces, which are then analyzed using molecular genetic techniques in order to determine individual genotypes with respect to a usually small number of loci. Two types of classification error can arise in such analyses. First, if only a small number of loci is examined, then there may be nonnegligible probabilities that multiple individual animals will have the same genotypes. The second type of error arises during the polymerase chain reaction (PCR) process and can result from failure of alleles to amplify (allelic dropout) or from PCR inhibitors in hair and feces that produce the appearance of false alleles or misprinting (Creel et al., 2003). Lukacs and Burnham developed models that formally incorporate possible misclassification of samples resulting from these errors. These models permit estimation of parameters such as abundance and survival in a manner that properly incorporates this uncertainty of individual identity. We anticipate that noninvasive sampling based on molecular genetic analyses of hair or feces will become extremely important for some species, and that the models of Lukacs and Burnham will become very popular for such analyses. MacKenzie & Nichols (2004) discuss the use of occupancy (proportion of patches or habitat area that is occupied) as a surrogate for abundance. In cases of territorial species and where birds occur at low densities, the number of occupied patches may provide a reasonable estimate of abundance. In other cases, occupancy can be viewed as providing information about one tail of the abundance distribution, P (N = 0). The motivation for considering occupancy as a surrogate for abundance is that occupancy is based on so–called presence–absence surveys that are frequently less expensive of time and effort than methods that estimate abundance directly. We describe one set of models that can be used to estimate occupancy for a single season and another that can be used to estimate parameters such as local probabilities of extinction and colonization that are associated with occupancy dynamics. We outline a possible hybrid approach that combines occupancy data with data on marked individuals in order to betterexplore the mechanisms underlying occupancy dynamics. These five presentations made for an interesting session containing useful information and recommendations for future work. A number of themes connecting these presentations could be emphasized. For example, two of the presentations considered alternatives to standard capture–recapture sampling that can be used to draw inferences about abundance, or a portion of the abundance distribution, with field methods that should be less expensive than usual capture–recapture approaches of handling animals. We believe that the most important theme of the session was the emphasis on the processes responsible for changes in abundance. In particular, we are excited by the potential for using hierarchical models as a means of investigating relationships among vital rates and as a means of combining multiple sources of data relevant to system dynamics. Indeed, we expect the importance of this session theme to be reflected in the content and presentations of the next EURING meeting.

Animal Biodiversity and Conservation

Mechanisms for retention of low molecular weight organic carbon varies with soil depth at a coastal prairie ecosystem

Though primary sources of carbon (C) to soil are plant inputs (e.g., rhizodeposits), the role of microorganisms as mediators of soil organic carbon (SOC) retention is increasingly recognized. Yet, insufficient knowledge of sub-soil processes complicates attempts to describe microbial-driven C cycling at depth as most studies of microbial-mineral-C interactions focus on surface horizons. We leveraged a well-studied paleo-marine terrace (90 ka) located near Santa Cruz, CA, to characterize the short-term (days to weeks) and intermediate-term (months to years) fate of two low molecular weight organic carbon. compounds at three depths in the soil profile (∼25 cm, A horizon; ∼75 cm A/B transition; and ∼125 cm, B horizon). We employed isotopically-labeled glucose (GLU) and oxalic acid (OXA) to represent two common classes of rhizodeposits: carbohydrates and organic acids. Using a combination of laboratory (9 d) and field (490 d) incubations, we traced the fate of GLU-C and OXA-C through dissolved-, metal-associated-, and microbially-respired CO2 and bulk SOC pools. Our results suggest new SOC retention (i.e., defined as 13C label identified in solid or aqueous fractions) over intermediate time frames (490 d) is correlated with patterns in short-term (9 d) cycling dynamics, which in turn is related to the theoretical efficiency by which microorganisms process each substrate. For all horizons (A, A/B, and B) GLU-C was converted to CO2 more quickly than OXA-C with modeled decomposition rates ∼2–4 times faster for GLU depending on microbial density (higher in A than B horizon). The faster decomposition rates of GLU-C increased fractional recovery (0.399 ± 0.026 to 0.504 ± 0.030 for GLU-C) compared to OXA-C (0.035 ± 0.003 to 0.127 ± 0.010) among all horizons in our field experiment (490 d). Though the overall proportion of GLU-C recovered in solid fractions did not vary significantly with horizon, based on 13C recovered in aqueous fractions the apparent mechanism for retention did. After the 9-d laboratory incubation, fractional recovery for GLU-C among C pools associated with microbial biomass was almost 20× higher than OXA-C (0.192 versus 0.010, respectively across all horizons). More than a year later, 43–46% of GLU-C retained in the field incubation was extractable with a neutral salt (representing a pool of soil C residing within or available to microbial biomass) among A and A/B horizons, while only 6% of retained GLU-C was similarly extractable in the B horizon. Thus, it appears among depths with higher microbial density (A, A/B horizons), anabolic recycling is the most likely process contributing to the persistence of glucose C, whereas abiotic sinks contributed more to intermediate-term stability for GLU-C in the B horizon. By contrast, most OXA-C was lost, presumably as CO2, over the short-term from the A and A/B horizons (fractional recovery: 0.136 ± 0.011 and 0.091 ± 0.002, respectively). However, though substantially lower than GLU-C recovered at the conclusion of our field experiment, the fraction of oxalic acid C retained in the B horizon over both short- (0.72 ± 0.037) and intermediate-time (0.127 ± 0.010) frames was several-fold higher than for overlying horizons. The specific process(es) (e.g., more efficient microbial utilization, metal-organic complexation, direct adsorption to the mineral matrix, etc.) contributing to higher retention for OXA-C at depth are discussed but remain unresolved.

Soil Biology and Biochemistry