These alterations directly increased the rate of biliary sterol excretion and increased

uptake of LDL cholesterol by the liver via the up-regulation of LDL-R. This study was supported by the National Council of Scientific and Technological Development (CNPq, Brazil; CNPq No. 480068/2009-7). We thank Maria Terezinha Bahia (Chagas’ Disease Laboratory, Federal University of Ouro Preto) for the use of the real-time PCR ABI 7300 equipment (Applied Biosystems). M.O.S and M.L.P were sponsored by a fellowship from CAPES and CNPq, respectively. We are also grateful to Rinaldo Cardoso dos Santos for his suggestions and careful review of the manuscript. “
“There has been an error with regard to Fig. 1. The orientation Screening Library mw of ICP gene cassette is given from EcoRI to HindIII where it should be from HindIII to EcoRI. This error is deeply regretted. The correct map of T-DNA is given below. “
“Acerola (Malpighia emarginata D.C.), also known as Barbados Cherry, is a tropical

fruit of great economic and nutritional value because of its high content of vitamin C, which is associated with the presence of carotenoids, anthocyanins, iron and calcium. Acerola’s consumption in natura is limited because it is a small fruit with relatively large seeds and is very perishable. The fruit, however, has a good pulp yield, facilitating the development of several IDH phosphorylation industrial products. Acerola has been processed in the form of juices, jams, ice creams, syrups, liqueurs and fruit syrups, among other products ( Soares Filho & Oliveira, 2003). In this context, processed products, such as frozen pulp and concentrated pulp, have economic importance; pulp production is a profitable activity that allows the freshly harvested perishable fruit to be stored and reprocessed off-season. The preservation of nutritional constituents during processing represents a major challenge for the traditional

techniques of pulp production. Processing generally involves heat treatment that can reduce the nutritional and organoleptic quality of the product. Over the years, new processing technologies have emerged to reduce or to eliminate the exposure ifenprodil of the fruit to heat. Ohmic heating is one alternative pulp pasteurization process. This technology can provide rapid and uniform heating, resulting in less thermal damage to labile substances such as vitamins and pigments (Castro et al., 2004 and Sarang et al., 2008). Ohmic heating is defined as a process where electric currents pass through foods to heat them by internally generated energy, without involving any heating medium or heat transfer surface (Castro, Teixeira, Salengke, Sastry, & Vicente, 2003). This heating technology is particularly interesting for viscous products and foods containing particulates because it simultaneously generates heat in both phases and does not need to transfer heat either through a solid–liquid interface or within a solid (de Alwis and Fryer, 1990, Imai et al.

Dopływy wspólnej żyły płucnej są stopniowo włączane w obręb przyszłego lewego przedsionka, co powoduje, że tworzą one jego tylną ścianę. Odmienne pochodzenie przedsionków zarówno pod względem tworzących je populacji komórek, jak i ekspresji genów, powoduje różnice selleck screening library morfologiczne tych jam opisane w dalszej części opracowania. Wspólny przedsionek jest charakterystyczny tylko dla wczesnego

rozwoju zarodkowego. Następnie od góry i tyłu dochodzi do wrastania fałdu tworzącego przegrodę pierwotną, zwaną inaczej pierwszą (Ryc. 4). Nie dochodzi ona jednak nigdy do poziomu kanału przedsionkowo-komorowego, ale pozostawia u dołu charakterystyczny otwór zwany pierwotnym (pierwszym). Jego zamknięcie następuje w czasie tworzenia się zastawek przedsionkowo-komorowych przy udziale poduszeczek wsierdziowych kanału przedsionkowo-komorowego i kolca przedsionkowego zamykającego go ostatecznie około 5. tygodnia rozwoju [19, 20]. Wspomniany kolec przedsionkowy odgrywa w tym procesie kluczową rolę, bowiem jego deficyt jest znanym już czynnikiem powodującym powstawanie wrodzonej wady serca pod postacią całkowitego ubytku przegrody przedsionkowo-komorowej (zwanego dawniej wspólnym kanałem przedsionkowo-komorowym) selleck u płodów z trisomią 21. chromosomu (zespołem Downa), malformacji znamiennie częściej

występującej u osób z tą aberracją [21]. Podczas gdy otwór pierwotny nie uległ jeszcze zamknięciu, w górnej części przegrody pierwotnej tworzą się fenestracje, które zespalając się ze sobą, uformują otwór wtórny (drugi). Fałd wpuklający się od góry pomiędzy żyłą główną górną a żyłami płucnymi utworzy przegrodę wtórną (drugą). Otwór drugi pełni u płodu niezwykle istotną funkcję, bowiem pozwala na swobodny przepływ krwi z żyły głównej dolnej (a co za tym idzie – utlenowanej krwi z łożyska Amylase doprowadzonej drogą żyły pępkowej) przez prawy przedsionek do przedsionka lewego, z częściowym ominięciem prawej części

serca [22]. Po urodzeniu otwór drugi jest zamykany przez przegrodę pierwotną (zastawkę otworu owalnego). Jeżeli proces ten ulega zaburzeniu, dochodzi do powstania po urodzeniu ubytku przegrody międzyprzedsionkowej typu otworu drugiego (atrial septal defect ostrium secundum type; ASD II), gdy przegroda pierwotna jest zbyt mała w stosunku do otworu drugiego, lub też przetrwałego otworu owalnego (patent foramen ovale; PFO), gdy nie dochodzi do całkowitego zrośnięcia przegrody pierwotnej i wtórnej [3, 23]. Wydawać by się mogło, iż proces tworzenia przegrody międzyprzedsionkowej może w warunkach nieprawidłowych skutkować wyłącznie powstaniem ubytku w jej obrębie. Należy jednak zwrócić szczególną uwagę na miejsce, w którym dochodzi do wpuklania się owej przegrody.

Rocket accessions were selected from three European gene banks based upon information provided by Elsoms Seeds Ltd. (Spalding, Lincolnshire, UK). In total 19 were sourced; 2 from the Centre for Genetic Resources in the Netherlands (CGN, Wageningen, The Netherlands), 12 from the Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung (IPK, Gatersleben, Germany), and 5 from the University of Warwick Crop Centre Genetic Resources Unit (Wellesbourne, UK; formerly Warwick HRI). A further

16 commercial varieties were collected: 13 were independently sourced from retailers, 1 provided by Elsoms Seeds Ltd., and 2 from Bakkavor Group Ltd. (Bourne, Lincolnshire, UK). Three biological replicates of each accession/variety Z-VAD-FMK solubility dmso were germinated under controlled environmental conditions (in Saxcil growth cabinets) after being sown in a random sequence. Long-day Roxadustat purchase lighting was used (16 h light, 8 h dark) at an intensity of 200 μmol m−2 s−1 (equivalent to 10,800 Lux of sunlight). Daytime temperatures were set at 20 °C and nighttime temperatures at 14 °C. Seedlings were grown for ten days in seedling trays and then transplanted to larger trays; four plants of each replicate were grown on. Plants were grown for another twenty days and then leaves from the four plants were harvested together. Sampling for each plant took approximately one minute from the cutting of the leaves

at the petiole to being placed in zip-loc freezer bags on dry ice inside a polystyrene container (with lid). For health and safety reasons it was decided that liquid nitrogen would not be used in this process. Thirty days was chosen as the optimum point of harvest as it reflects the typical number of days commercial growers grow their crop after sowing. Bags were placed in a −80 °C freezer immediately after harvest and transport was completed (<30 min). Samples were freeze-dried

Pregnenolone in batches for three days (in a Vertis Bench-top Series). Leaves from each rep were ground into a fine powder using a combination of pestle and mortar and miniature coffee grinder (De’Longhi KG49, Treviso, Italy). All solvents and chemicals used were of LC–MS grade and obtained from Sigma–Aldrich (Poole, UK) unless otherwise stated. The following method was adapted from Pasini, Verardo, Caboni, and D’Antuono (2012), Jin et al. (2009). Three experimental replicates of each biological rep were prepared as follows: 40 mg of ground rocket powder was heated in a dry-block at 75 °C for 2 min, as suggested by Pasini, Verardo, Caboni, and D’Antuono (2012), as a precautionary measure to inactivate as much myrosinase enzyme as possible before liquid extraction. 1 ml of preheated 70% (v/v) methanol (70 °C) was then added to each sample and placed in a water bath for 20 min at 70 °C. Samples were then centrifuged for 5 min (6000 rpm, 18 °C) to collect loose material into a pellet. The supernatant was then taken and put into fresh Eppendorf tubes.

For example, proteins that possess the ellipsoidal shape two types of orientation of adsorbed molecules may occur: side- and end-on adsorption if major axis is parallel and perpendicular to solid surface, respectively [23]. Conformational change or change in lateral interaction may run concurrently with the adsorption process. Discontinuities in the adsorption isotherms as observed in this work may indicate different concentration-dependent orientation of adsorbed click here molecules at interface due to a non-spherical protein shape [24]. The ability of BSA to establish strong molecule-molecule interaction when in contact

with HA surface was confirmed by AFM analyses performed in sintered discs (1 cm2 in diameter) before and after the protein adsorption. For BSA initial concentration of 0.05 mg/mL with 24 hours incubation time, the AFM images revealed a thin BSA film covering the HA disc surface still maintaining visible the HA grain boundaries (Fig. 3b). The protein was homogenously distributed over the HA disc surface but small aggregates of protein could be detected at grains surface. The adsorption

pattern changed drastically when higher BSA concentrations were used (2.0 mg/mL). In that case, the adsorption was not homogeneous and large aggregates of protein were formed in different parts of discs surface as shown in Fig. 3d and f. These agglomerates can be produced when surface coverage exceeds a saturation value as suggested by Xu et al. [14]. The reactivity of HA + BSA surface was evaluated by Selleckchem TSA HDAC its capability to induce the crystallization of a new calcium phosphate when the surface was in contact with a simulating body fluid solution (n-SBF). For this evaluation, HA discs were coated Rutecarpine with a high BSA concentration (1.42 × 10−6 mmol/cm2) in order to cover the whole HA surface with protein layers and to promote a strong protein–protein interaction on HA surface. In such condition the stability of BSA film on

HA surface was assured for periods up to 7 days. After 7 days soaking in n-SBF, HA discs with BSA (HA + BSA/SBF) and without BSA (HA/SBF) were both fully covered by a thick layer of crystalline particles as observed by SEM, Fig. 4a and b. That thick layer was further characterized as a poorly crystalline calcium phosphate phase (CaP) by FTIRM-ATR and GIXRD measurements. The new layer presented similar crystal morphology in discs with and without BSA. In order to investigate the influence of BSA on the formation of the apatite layer the precipitation rate of calcium and phosphorous onto discs surface were followed by ICP. As shown in Fig. 5, calcium and phosphorous concentration on n-SBF solution decreased gradually with time confirming the precipitation of a new calcium phosphate on HA and HA + BSA discs surface.

Whereas the action execution is an obvious extension of inner intentions in response to specific stimuli, the “primum movens” of our knowledge, selleck i.e. the link between action performance and conscious perception of causal agency, remains intriguing. From the discussion above, we may infer that a personal identity is

psychologically installed in the agent’s mind in order to observe autopoiesis: achieving the goal of self-organisation. In the overall picture, “Free Will” does not exist: it is only a belief of the inner observer. However, provided the inner observer survives, this illusion is justified since it is like an energy gear for such a cognitive system: it makes PI imagination work harder and better, i.e. it is the basic requirement for the reward circuitry operating at maximal efficiency; otherwise, according to Maturana and Varela (1980) and Varela, Thompson, and Rosch (1991) the system would disintegrate (Bignetti, 2001, Bignetti, 2003 and Bignetti, 2004). Cognitive systems do not operate by representing world as a sum of independent components; knowledge is enacted as a series

of distinct elements, inseparable from structures embodied by cognitive systems. On the one hand, the term “enaction” emphasises the growing conviction that cognition is not the representation of a pregiven buy LBH589 world by a pregiven mind but is rather the enactment of a world on the basis of its history and the variety of actions that a being in the world performs; on the other hand, “embodiment” provides a systemic and dynamic framework for understanding how a cognitive self (a mind) can arise in an organism in the midst of its operational cycles of internal regulation and ongoing sensorimotor coupling. Another paper solely devoted Isotretinoin to discussing the fundamentals of TBM in connection with the stimulating thought of Varela would be useful. TBM argues that ‘free’ decisions are determined by early brain activity. Libet’s pioneering and controversial studies (Libet, 1983 and Libet, 2004) on the timing of action

decisions taken in the brain, observed the onset of early electrical activity, known as the “readiness potential” (RP), prior to the onset of conscious will. More recently, it has been shown that the outcome of a decision can be encoded in the brain activity of the prefrontal and parietal cortex up to 10 s before it enters our awareness. This delay presumably reflects the operation of a network of higher level control areas that begin to prepare an upcoming decision long before it enters our awareness (Soon, Brass, Heinze, & Haynes, 2008). This data is even more striking in the light of other research suggesting that the decision to move, and possibly the ability to halt that movement at the last second, may be the result of unconscious processing (Matsuhashi & Hallett, 2008).

Genetic diversity estimates across loci indicated that ISS did not reduce mean allele

indices either in the natural regeneration of the managed stand or in the managed stand itself when compared to the old growth forest. Across loci, 12 out of 119 alleles were lost in the succeeding generation in the managed stand and 16 out of 123 in the old growth stand. In contrast, saplings from the old growth were more successful in recruiting new alleles into their population; they recruited 15 alleles not present in the sampled adult cohort in comparison to the managed stand where the saplings recruited 9 new alleles. All alleles lost in the next generation but one from the old growth were rare alleles. Majority of newly recruited alleles were also rare; 7 in the managed and 14 in the old growth stands. PLX3397 mouse The inbreeding coefficient FIS significantly departed from the expected value PD0332991 molecular weight only in the sapling population in the old growth forest (FIS = 0.052, p = 0.017) because of the departures from the expected value at locus Fs3 (FIS = 0.229, p = 0.021). This was most likely caused by the presence of null alleles at this locus as identified with the Micro-Checker programme. Null alleles were also detected at loci Fs10 and Fs15 in the adult phase of the old growth stand but global FIS for this cohort did not significantly depart from the expected value under random mating (FIS = 0.016, p = 0.270). The lack of inbreeding in the study was anticipated

as inbreeding was not expected to occur in an outcrossing species like beech. Temporal changes in allele frequencies that could not be attributed

only to genetic drift and sampling error between the cohorts were detected in both the managed and old growth stands (Table 2, Fig. 2). In the managed stand significant temporal changes in allele frequencies were detected at loci Fs5, Fs6 and Fs8 while in the old growth temporal changes caused by factors other than genetic drift, sampling error and management were observed at loci Fs6 and Fs10. Repeating the simulations with frequencies adjusted for null alleles, according to Chakraborty et al., 1992 and Van Oosterhout et al., 2004, that were implemented in the Micro-Checker programme for loci exhibiting null alleles (Fs3, Fs10 and Fs15), changed the observed FST values but did not alter the rejection of the null hypothesis Thiamet G for locus Fs10 and did not result in its rejection for the other two loci. FST values did not significantly differ from the expected values for any of the loci either in the managed or old growth stands after applying Bonferroni corrections for multiple comparisons. However, before the application of correction for multiple comparisons, p values for loci Fs5 and Fs6 in the managed stand and loci Fs6 and Fs10 in the old growth stand were lower than 0.05, indicating a good fit with the results obtained with the FT, ST and WT tests. FST value between adults and saplings in the managed stand (0.0042, p = 0.

[pause] We’re all in similar situations. It’s like some person who’s blindfolded and in the middle of a field [continues with the entire metaphor about falling in hole (representing emotion) and trying to dig out (representing attempts to regulate ABT-263 chemical structure emotion)]… And sometimes we can’t tell that our shovels aren’t working because we’re digging so hard, and we want it to work. I think you’ve been trying the logical thing. If you have emotions you don’t like, you try to get rid of them or push them away. And it’s supposed to work, right? But

our experience tells us something different. So, maybe the first step can be to stop digging and drop the shovel. If you’re the therapist and you tell someone that the first step is

to let go, how do you think they’re going to respond? During these sessions, participants assessed the different ways in which they had tried to “dig” their way free from difficult thoughts and feelings and how effective those strategies had been. Both participants identified binge eating as strategies they used to distract themselves from or avoid unpleasant internal events. In addition to not being able to fully eliminate unwanted thoughts and feelings, participants often experienced feelings of guilt, shame, sadness, self-loathing, and frustration after binge eating. Once the participants became aware of the futility of efforts www.selleckchem.com/mTOR.html to control unwanted internal events, the next step was to teach acceptance and mindfulness skills (e.g., increased awareness of and contact with internal events as they are, fully, without making efforts to eliminate them) as behavioral alternatives to control efforts. Beginning with MycoClean Mycoplasma Removal Kit the first session, the therapist introduced a series of brief mindfulness exercises in order to build the skill of gently and nonjudgmentally paying attention to specific objects or internal experiences as they are without trying to alter or get rid of them (Kabat-Zinn, 1990). For example, in a brief

mindfulness exercise, participants intentionally monitored physiological sensations and/or the act of breathing for 1 or 2 minutes. During the exercise, the participants were instructed to notice how their attention drifted away from breathing and other physical sensations and to bring their focus back to the present moment when they noticed that their attention had drifted away. In one particular exercise, participants also practiced a mindful eating exercise using a raisin (Safer, Telch, & Chen, 2009, pp.102–103), which was based on an exercise described by Kabat-Zinn (1990). The purpose of the mindful eating exercise was to help participants increase their awareness in the context of eating. Increased awareness was particularly important because the behavior of eating often evoked intense unwanted emotions and thoughts. In this exercise, the participants were first asked to notice what emotional and/or situational triggers often preceded binge eating.

WNV can cause poliomyelitis-like illness or acute flaccid paralysis in WNV-infected

persons, which is histologically confirmed in the grey matter of the anterior spinal cord and in the brainstem of postmortem tissues (Doron et al., 2003, Fratkin et al., 2004, Jeha et al., 2003, Sejvar Lumacaftor price et al., 2005 and Sejvar et al., 2003b). Similar histopathology occurs in WNV-infected hamsters (Morrey et al., 2008b, Samuel et al., 2007, Siddharthan et al., 2009 and Xiao et al., 2001) and mice (Hunsperger and Roehrig, 2006) where the ventral cord has lymphocytic infiltration, perivascular cuffing, and neurophagia. Similar signs are documented with nearly all flavivirus encephalitides, i.e., Japanese encephalitis virus (JEV) (Johnson, 1987), tick-borne encephalitis (TBE) virus (Gelpi et al., 2005), and the murine Modoc virus (Leyssen et al., 2003). Observing histopathological changes in the central nervous system (CNS), however, does not necessarily cause or indicate the types of neurological deficits. For example, the spinal cord functions are vast and diverse, where the cord acts as a conduit for descending motor functions, as a conduit for ascending

sensory information, and as a center for coordinating sensory/motor reflexes. Essentially, it is a conduit between the brain and nearly all other body functions. Therefore, histopathological damage to the spinal cord by WNV could affect a wide range of neurological disease phenotypes. Since WNV clearly causes motor function deficits in selleck chemicals llc human subjects, human clinical procedures employed for evaluating WNND and other motor diseases have been adapted for measurement of motor functions in rodents infected with WNV. In neurodegenerative diseases such as poliomyelitis (Ohka and Nomoto, 2001) and amyotrophic lateral sclerosis (Rashidipour and Chan, 2008 and Shefner et al., 2006), the loss of motor neurons can be clinically detected by using electrophysiological motor unit

number estimation (MUNE) (Dantes and McComas, 1991), where a motor unit consists of a motor neuron and all its associated muscle fibers. Since a presumptive use of MUNE in the Rucaparib cell line human WNV infection appears to be a possible marker for muscle weakness and clinical recovery (Cao et al., 2005), the MUNE procedure was adapted for use in hamsters (Siddharthan et al., 2009). To perform the MUNE procedure, the rostral sciatic nerve is stimulated with incremental increases of voltage. The resulting M-wave depolarization and polarization voltages are recorded at the plantar aspect of the hind limb. As the stimulus is increased, more motor units are recruited or activated. The increased activation of motor units is detected by incremental jumps in the amplitude of the M-wave. The more incremental jumps that are detected, the more motor units the animal possesses.

, 2009, Edsall et al., 1988 and Leach, 1991) but commercial harvest is now heavily restricted and recreational catch of four major sport fishes (walleye, yellow

perch, smallmouth bass and muskellunge) is a more common activity ( Thomas and Haas, 2004). The fish community of LSC has been diverse and abundant with about 70 species of warm and cool-water species, including yellow perch, walleye, smallmouth bass (Micropterus dolomieui) and muskellunge as well as introduced species such as round gobies ( Leach, 1991 and Thomas and Haas, 2004). The wetland area of LSC was much greater historically than at present (especially along the Michigan side). It is estimated that 72% of the wetland learn more area was lost from 1873 to 1973 mainly due to urbanization (Jaworski and Raphael, 1976 and Leach, 1991). Conversion of wetlands to agriculture

was also common on the Ontario side. Emergent wetland vegetation, including cattails (Typha latifolia, Typha angustifolia), bulrush (Schoenoplectus tabernaemontani), common reed (Phragmites australis) and spike rush (Eleocharis quadrangulata) were common in undeveloped areas including the St. Clair Flats and the eastern shoreline ( Edsall et al., 1988 and Leach, 1991). For migratory birds like mallards, Ribociclib clinical trial black ducks, Canada geese and tundra swans, the vast wetlands provided essential flyway resting and feeding habitat ( Leach, 1991). Most of the native fish species spawned along the St. Clair Flats or along the Rutecarpine shoreline areas adjacent to the tributaries ( Goodyear et al., 1982 and Leach, 1991). The invasive common reed (P. australis) expanded across LSC when low lake levels followed the high lake levels in1986. P. australis can now be found along the coast line of LSC and poses problems because it forms thick strands, reduces functionality, biodiversity, and property values ( USGS Great Lakes Science Center, 2011 and Wilcox, 2012). Once Phragmites is established it can be difficult and expensive to remove

( USGS Great Lakes Science Center, 2011). In summary, the natural system of LSC has been influenced by human activities (i.e. contaminants and spread of invasive species), but the ecological condition also influences humans that depend on it for drinking water, recreational activities, and fishing. Thus identifying these components and linkages between human and natural systems is critical in planning for sustainability. The ecological condition and ecosystem services of LSC depend to a great extent on the human population, land use, climate and technological advances in water and wastewater management. We identified three periods during the last century that indicate fundamental changes to the socioeconomic system that might be appropriate for understanding changes to the ecology of LSC (Table 1).

9). Depth-averaged sand percentages fall between 74 and 92% for all samples analyzed; core-averaged organic matter percentages are between 1.5 and 2.4, respectively (Fig. 9). As cores show an overall low degree of grain-size variance with depth, likely attributed to a very high degree of bioturbation within the pond, depth-averaged percentages of organic matter were utilized in the construction of the pond-wide

correction factor for isolating the clastic sediment component (Co; Table 2 and Fig. 8). Maps of the 1974 and 2012 pond floor show sedimentation has most heavily affected the shore-proximal selleck screening library parts of the pond ( Fig. 7). An isopach map of post-1974 sediment thickness shows accretion of up to 1.5 m in select nearshore areas, which thins to the NE part of the basin, where only 0–25 cm of positive elevation change are recorded ( Fig. 7C and D). The total volume of post-1974 sediment in the pond approximates 6228 m3 based on the data. A dry inorganic sediment

weight is calculated from this measured sediment volume by applying factors for core compaction (Cc), organic sediment fraction (Co), and volume-to-weight conversion (Cvw) as shown in Fig. 8. Fig. 10 shows the spatial distribution of values for each of these conversion/correction factors used. Using this approach of spatial integration of correction values SB203580 chemical structure the calculated weight of inorganic sediment in Lily Pond sequestered since 1974 approximates 4,825,618 kg; this number decreases to 807,330 kg applying the lowest correction/conversion values as a spatial constant and 10,083,331 kg using the highest ( Table 3), providing an error envelope based on empirical data. All USLE factors used in the model are assumed to be well-constrained with the exception of the C-factor. Land managers interested GBA3 in developing similar USLE models for their particular regions of interest

would face the same dilemma given that data on soil, climate, and topographic variables are more easily accessed than detailed land-cover data. K-factors generally do not vary by an excessive range as do C-factors, which can show a very high degree of spatial and temporal variance; soils within the study area, for example, are comparable in their textural and compositional characteristics and therefore have similar K-values ( Lessig et al., 1971). The R-factor varies tremendously over the short-term (at the event-scale); however, the USLE operates on a long-term basis and applies an empirically constrained, time-averaged R-value, which varies little over large spatial scales ( Wischmeier and Smith, 1965). The SL-factor is invariable over time and tightly constrained from digital terrain analyses using a USGS 3 m DEM. The C-factor, shown to exert the single strongest control on soil-erosion model variance ( Toy et al., 1999), remains an unconstrained factor.