, 2005a, Erlandson et al., 2005b and Rick et al., 2008a). By 7000 years ago, the Chumash also appear to have introduced dogs and foxes to the island, which further affected the terrestrial ecology (Rick et al., 2008b, Rick et al., 2009a and Rick et al., 2009b). Millions of shellfish were harvested from island waters annually and signatures of this intensive predation have been

documented in the declining size of mussel, abalone, and limpet shells in island middens beginning as much as 7000 years ago (Fig. 5; Erlandson et al., 2009, Erlandson et al., 2011a and Erlandson et al., 2011b). Studies of pinniped remains from island middens also show that the abundance of northern elephant seals (Mirounga angustirostris) Angiogenesis inhibitor and Guadalupe fur seals (Arctocephalus townsendi) is very different today than the rest of the Holocene, probably due to the combined effects of ancient subsistence hunting and historic commercial seal hunting ( Braje et al., 2011 and Rick et al., 2011). In summary, although California’s Channel Islands are often

considered to be pristine and natural ecosystems recovering from recent ranching and overfishing, they have been shaped by more than 12,000 years of human activity. It has taken decades of intensive archeological buy GDC-0941 and paleoecological research to document this deep anthropogenic history. As other coastal areas around the world are studied, similar stories of long-term human alteration on islands and coastlines are emerging (e.g., Anderson, 2008, Kirch, 2005, Rick and Erlandson, 2008, Rick et al., 2013a and Rick et al., 2013b). Worldwide, long shell midden sequences provide distinctive stratigraphic markers for ancient and widespread human presence in coastal and other aquatic landscapes, as well as the profound effects humans have had on them. In coastal, riverine, and lacustrine settings around the world, there is a signature of intensive human exploitation of coastal and other aquatic ecosystems that satisfies the requirements of a stratigraphic

marker for the Anthropocene. This signature can be clearly seen geologically and archeologically in the widespread appearance between HSP90 about 12,000 and 6000 years ago of anthropogenic shell midden soils that are as (or more) dramatic as the plaggen soils of Europe or the terra preta soils of the Amazon (e.g., Blume and Leinweber, 2004, Certini and Scalenghe, 2011, Schmidt et al., 2013 and Simpson et al., 1998). Similar to these other anthropogenic soils, the creation of shell middens often contributes to distinctive soil conditions that support unique plant communities and other visible components of an anthropogenic ecosystem. When combined with other anthropogenic soil types created by early agricultural communities in Africa, Eurasia, the Americas, and many Pacific Islands, shell middens are potentially powerful stratigraphic markers documenting the widespread ecological transformations caused by prehistoric humans around the world.

Hillslope failure, river channel widening, and/or construction activity may mobilize sediment from deeper (i.e., meters) sources. Aeolian deposition may be a third source, although

no evidence supports aeolian deposition as a significant source to the rivers studied here. The relative contributions from these sources may change both temporally and spatially in a river. These changes allow only limited AZD2281 manufacturer conclusions to be drawn from a single data point, limiting the success of a mitigation effort that is applied uniformly across a watershed. Contemporary sediment sources are frequently augmented and supplemented by legacy sediment. Legacy sediment comes from anthropogenic sources and activities, such as disturbances in land use/cover and/or surficial processes (James, 2013). For rivers, legacy sediments can originate from incised floodplains (Walter and Merritts, 2008), impoundments behind dams (Merritts et al., 2011), increased hillslope erosion due to historic deforestation (DeRose et al., 1993 and Jennings et al., 2003), and anthropogenic activities

such as construction selleck inhibitor and land use changes (Wolman and Schick, 1967 and Croke et al., 2001). Legacy sediment can also deliver high loads of contaminants to river systems (Cave et al., 2005 and Lecce et al., 2008). The current supply of sediment is high (Hooke, 2000), as humans are one of the greatest current geomorphic agents. Consequently, combining legacy sediment with increased anthropogenic geomorphic activity makes it even more important to identify the source of sediments in rivers. Sediment sources can be distinguished NADPH-cytochrome-c2 reductase using the radionuclides lead-210 (210Pb) and cesium-137 (137Cs). 210Pb is a naturally-occurring isotope resulting from the decay of 238Uranium in rock to eventually 222Radon. This gas diffuses into the atmosphere and decays into excess 210Pb, which eventually settles to the ground. This diffusion process creates a fairly consistent level of excess 210Pb in

the atmosphere and minimizes local differences that exist in the production of radon. Rain and settling can subsequently result in the deposition of excess 210Pb, with a half-life of 22.3 years. This atmospheric deposition of excess 210Pb, is added to the background levels that originate from the decay of radon in the soil. “Excess” atmospheric 210Pb occurs because, if the material (in this case the sediment) is isolated from the source (i.e., the atmosphere), this level will decay and decrease in activity. As this excess 210Pb is then correlated with the time of surficial exposure, it is commonly used as a sediment tracer (e.g., D’Haen et al., 2012, Foster et al., 2007, Whiting et al., 2005 and Matisoff et al., 2002). 137Cs is also used as a sediment tracer, although its source is different. It is the byproduct of nuclear fission through reactors and weapon activities, and is not naturally found in the world.

The upper bands were comparatively larger than expected and were revealed to be artifacts in PCR amplification (Band-C and Band-D in Fig. 1). Multiple band artifacts are common in the amplification of SSR sequences and can generally be removed by modifying the PCR conditions or the number of cycles [22], [23] and [24]. buy Erastin However, the examples described here are fundamentally different from the previous reports for two reasons: firstly, the larger bands could not be abolished by altering the PCR conditions; and, secondly, the reamplification of the larger bands showed the same band patterns as that of preliminary

PCR amplification. Based on all these findings together, it appears that the artifact bands are derived from heteroduplexes created by the combining and interruption between coexisting different amplicons. The appearance of multiple bands in PCR products has been regarded as one of the more serious obstacles to marker development and genetic mapping for recently duplicated plant genomes such as rapeseed (Brassica napus) [25] and P. ginseng [9] because they hinder genotyping against the mapping population as well as the authentication of cultivars. In

this study, a clear single band was successfully amplified by using a locus-specific primer designed on the basis of sequence variation between the two paralogous loci. The locus-specific primer was based on the SNP sequence of the polymorphic band of the gm47n marker. In addition to the SNP, T/C in Selleckchem Osimertinib Band-B and Band-A, we added another modified nucleotide, “G” instead of “A”, that resulted in a clear single band of PCR product in ginseng, as suggested by a previous report [17]. The clear single band was polymorphic between two cultivars, Chunpoong and Yunpoong, and segregated with a Mendelian single gene pattern in their F2 population ( Fig. 4). These results support our assumption that Band-A and Band-B are not heterozygous

forms, but instead are derived from different loci created by the recent genome duplication of P. ginseng [7]. Our method can be applied to other markers to overcome the genotyping difficulty caused by multiple bands in P. ginseng. Most plant species have undergone a few rounds of genome duplication [26] and [27]. We suggest Cepharanthine that this approach should be considered as an efficient method to avoid the misinterpretation of multiple band appearances in genome research on wild plant resources that may have undergone recent genome duplication. Utilization of upcoming ginseng genome sequence information will be a powerful tool for the development of indisputable and reliable markers and genetic mapping in P. ginseng. We are conducting whole-genome sequencing for the cultivar Chunpoong using the Illumina platform [28] and have identified many long paralogous genome sequence pairs from the draft sequence assembly. Each of the paralogous sequences can be mapped by developing paralogous locus-specific markers as suggested in this study.

XO generates ROS during the oxidation of hypoxanthine or xanthine [32], and Ohta et al [33] suggested

that the xanthine–XO system in the gastric mucosal tissue participates in the progression of gastric mucosal lesion. In the present study, increased MPO activity—an index of neutrophil infiltration—of the gastric lesion control group was reduced, and ROS-related parameters such as MDA content and XO activity were normalized by ginsenoside Re administration. From the present study, it seems likely that administration of ginsenoside Re exerts a preventive effect on the progression of C48/80-induced acute gastric mucosal lesions by protecting the gastric mucosal barrier and tissue against the attack of ROS derived from infiltrated neutrophils and the xanthine–XO system Lumacaftor datasheet through preservation of gastric mucus. The protein encoded by the Bcl2 gene is a regulator of programmed cell death and apoptosis. The cell survival-promoting activity of this protein is contrary to the cell death-promoting activity of Bax, a homologous protein that forms heterodimers with Bcl2 and accelerates rates of cell death [34]. The

expression of Bax is upregulated by the response of the cell to stress [35]. Bax protein significantly increased 3 h after hypoxic–ischemic brain injury in neonatal brain tissue [36] and it increased in gastric mucosa after ischemia–reperfusion damage [37]. In the present results, the predominant increase of Bax expression was discovered after C48/80-induced acute gastritis. We have click here observed that the increased Bax expression by C48/80 treatment was attenuated when ginsenoside Re was administered. In contrast to Bax, Bcl2 expression decreased after C48/80 induced acute gastritis and ginsenoside Re attenuated the diminution. In Western blotting analysis, the Bax/Bcl2 ratio result also confirmed the protective effects of ginsenoside Re on C48/80-induced

acute gastritis. In conclusion, the results of the present study indicate that ginsenoside Re exerts a preventive effect on the progression of C48/80-induced acute gastric mucosal lesion in rats, possibly by inducing mucus secretion and attenuating enhanced neutrophil infiltration, inflammation, and oxidative stress in gastric mucosa. The authors declare BCKDHB no conflicts of interest. This study was funded by the program of the Kyung Hee University (Seoul, South Korea) for the young medical researcher in 2008 (KHU-20081252). “
“Of the primary energy sources in the human body (carbohydrates, proteins, and lipids), lipids are the most efficient type of energy storage (9 kcal/g) and are hence much more prevalent than carbohydrates or proteins as a form of storage [1]. This makes the process of lipid release a crucial component in understanding human energy metabolism and pathology.

Trace metals are also high in the upstream Le Fever Dam pool sediment ( Kasper, 2010 and Peck and

Kasper, 2013). The elevated trace metal content in the Gorge Dam sediment reflects anthropogenic activities in the watershed well beyond the adjacent power plant. During much of the Second Period the Cuyahoga River served as a convenient way to dispose of the wastes from BMS-354825 chemical structure many anthropogenic activities (Moloney et al., 2011). Magnetic susceptibility, a proxy for CCP particles, increases at about the times (1930, 1940, and 1960) the power plant was expanded (Fig. 8). All four trace metal concentrations decline in the 1930s, possibly as the result of decreased anthropogenic pollution activities during the Great Depression. Between 1930 and 1940 the population of Cuyahoga Falls remained the same (Fig. 9). From 1940 to 1960 both the Pb concentration and the Cuyahoga Falls population increase (Fig. 8 and Fig. 9). Activities such find more as construction, automobile traffic, industry, urbanization and suburbanization related to the growing population contributed to the poor sediment quality within the Gorge Dam pool. The Clean Air Act (1970), Clean Water Act (1972) and a growing environmental awareness greatly contributed to bringing the Second Period to an end (Fig. 8). Maximum use of leaded gasoline occurred in 1970 nationwide,

locally, urban lead sources peaked at various times throughout the 1970s (Callender and Van Metre, 1997). The Third Period (1978–2011) period is defined by mud having greatly reduced amounts

of CCP, declining trace metals, and low magnetic concentration (Fig. 8). Although the four trace metals begin this period above the PEC, all decline below the PEC toward the present day following a similar trend identified in nearby Summit Lake (Haney, 2004) and in other U.S. reservoirs (Callender and Van Metre, 1997). The Gorge Dam pool sediment record shows a steady decline in Pb concentrations starting in about 1985. The decline in trace metals Levetiracetam in this period is a response to the Clean Air Act (1970), the Clean Water Act (1972), and declining industrial activity in the watershed. Also, in 1988, the Cuyahoga River was put on the list of Areas of Concern to help improve water quality in the Lake Erie basin (Moloney et al., 2011). The effectiveness of these environmental regulations is evident, because the last identifiable CCP layer in the dam pool sediment dates to about 1978, even though the coal-fired power plant continued to produce electricity until 1991 (Whitman et al., 2010, p. 80). Unlike monitoring programs that may take years to generate a record of a stream’s sediment load variability, dam pool sediments can quickly provide such a record, when dated with a high-resolution method such as 210Pb dating. A sediment load record obtained from a dam pool allows one to assess the range of variability since the dam was installed.

, 2005a, Erlandson et al., 2005b and Rick et al., 2008a). By 7000 years ago, the Chumash also appear to have introduced dogs and foxes to the island, which further affected the terrestrial ecology (Rick et al., 2008b, Rick et al., 2009a and Rick et al., 2009b). Millions of shellfish were harvested from island waters annually and signatures of this intensive predation have been

documented in the declining size of mussel, abalone, and limpet shells in island middens beginning as much as 7000 years ago (Fig. 5; Erlandson et al., 2009, Erlandson et al., 2011a and Erlandson et al., 2011b). Studies of pinniped remains from island middens also show that the abundance of northern elephant seals (Mirounga angustirostris) BGB324 mouse and Guadalupe fur seals (Arctocephalus townsendi) is very different today than the rest of the Holocene, probably due to the combined effects of ancient subsistence hunting and historic commercial seal hunting ( Braje et al., 2011 and Rick et al., 2011). In summary, although California’s Channel Islands are often

considered to be pristine and natural ecosystems recovering from recent ranching and overfishing, they have been shaped by more than 12,000 years of human activity. It has taken decades of intensive archeological HDAC inhibitor and paleoecological research to document this deep anthropogenic history. As other coastal areas around the world are studied, similar stories of long-term human alteration on islands and coastlines are emerging (e.g., Anderson, 2008, Kirch, 2005, Rick and Erlandson, 2008, Rick et al., 2013a and Rick et al., 2013b). Worldwide, long shell midden sequences provide distinctive stratigraphic markers for ancient and widespread human presence in coastal and other aquatic landscapes, as well as the profound effects humans have had on them. In coastal, riverine, and lacustrine settings around the world, there is a signature of intensive human exploitation of coastal and other aquatic ecosystems that satisfies the requirements of a stratigraphic

marker for the Anthropocene. This signature can be clearly seen geologically and archeologically in the widespread appearance between Adenosine triphosphate about 12,000 and 6000 years ago of anthropogenic shell midden soils that are as (or more) dramatic as the plaggen soils of Europe or the terra preta soils of the Amazon (e.g., Blume and Leinweber, 2004, Certini and Scalenghe, 2011, Schmidt et al., 2013 and Simpson et al., 1998). Similar to these other anthropogenic soils, the creation of shell middens often contributes to distinctive soil conditions that support unique plant communities and other visible components of an anthropogenic ecosystem. When combined with other anthropogenic soil types created by early agricultural communities in Africa, Eurasia, the Americas, and many Pacific Islands, shell middens are potentially powerful stratigraphic markers documenting the widespread ecological transformations caused by prehistoric humans around the world.

The effective cation exchange capacity was calculated as a molar ratio of exchangeable Al (Ex-Al3+) to the sum of exchangeable Ca (Ex-Ca2+), exchangeable Mg2+, exchangeable sodium (Ex-Na+),

Ex-K+, and Ex-Al3+[15]. The Al saturation was calculated as Al/effective cation exchange capacity. The soils were also extracted using 0.1M Na-pyrophosphate (pH 10.0; soil ratio: extractant 1:100, with shaking for 16 h) for organic Al (Alp) [16]. The Al in the extract solution was measured in duplicates using an atomic absorption spectrometry equipped with graphite furnace Bosutinib atomizer (PerkinElmer Analyst 700; PerkinElmer Inc., Norwalk, CT, USA). The data were statistically evaluated using the Data click here Processing System 11.0 edition for Windows [17] (Zhejiang University, Hangzhou, China). Data are presented as the mean ± standard deviation. Analysis of correlation was performed with three replicates. Some studies have indicated that unbalanced cations and nutrition disorders have contributed to a decline in ginseng

garden soil conditions [1] and [18]. A measurement of the major cations was carried out seasonally. Both concentrations of Ex-Na+ and Ex-K+ stayed relatively constant without obvious spatial variation during 2009; however, they sharply increased in the 0–5 cm depth in the spring of 2010 (Fig. 1A–J). The exception was the decrease in both the Ex-Na+ and Ex-K+ in transplanted 1-yr-old ginseng soils in the spring, which might be driven by individual factors. The Ex-Ca2+ concentration showed a decrease within a 1-yr cycle of investigation (Fig. 1K–O). For transplanted 1-yr-old ginseng soils particularly, the Ex-Ca2+ concentration sharply decreased selleck chemicals llc in the three depths after the spring of 2009 (Fig. 1N). Although the Ex-Ca2+ concentrations in

the transplanted 2-yr-old ginseng soil were constant, a value of approximately 0.4 was the lowest of the detected Ex-Ca2+ concentration data (Fig. 1O). The exchangeable Mg2+ concentrations were kept relatively constant at the three soil depths for the different aged ginsengs within a 1-yr cycle (Fig. 1P–T). The NH4+ concentrations showed sharp decreases at all three depths from the spring of 2009 (Fig. 2A–E). The decrease was more remarkable in the summer and autumn. There were two obvious exceptions: the increase of NH4+ in the 0–5 cm layer for the 1- and 3-yr-old ginseng soils during the next spring (Fig. 2A,C), which might have been driven by individual factors. The surface (0–5 cm) NO3− concentration exhibited a remarkable increase in the summer and autumn, and then sharply decreased to the original level by the next spring (Fig. 2F–L). The NO3− concentrations in the 0–5-cm layer peaked in the autumn and were over 10-fold greater than those in the spring (Fig. 2F–L).

52 (C-14), 33.13 (C-15), 27.25 (C-16), 51.40 (C-17), 16.94 (C-18), 17.09 (C-19), 140.66 (C-20), 13.66 (C-21), 123.82 (C-22), 27.95 (C-23), 123.92 (C-24), 131.74 (C-25), 26.18 (C-26), 18.22 (C-27), 29.33 (C-28), 16.31 (C-29), 17.52 (C-30), 105.62 (3-Glc C-1′), 83.95 (3-Glc C-2′), 78.76 (3-Glc C-3′), 72.12 (3-Glc C-4′), 78.45 (3-Glc C-5′), 63.19 (3-Glc C-6′), 106.55 (3-Glc C-1″), see more 77.64 (3-Glc C-2″), 78.84 (3-Glc C-3″), 72.15 (3-Glc C-4″), 78.62 (3-Glc C-5″), 63.34 (3-Glc C-6″) (Fig. 2) [22]. MCF-7 (HER2-/ER+) and MDA-MB-453 (HER2+/ER–) human breast cancer cell lines

were maintained using RPMI 1640 medium supplemented with 10% (vol/vol) FBS (Welgene, Daegu, South Korea) plus 100 units/mL penicillin and streptomycin in a 5% carbon dioxide air incubator at 37°C. Cell cytotoxicity was measured by MTT assay. Cells were seeded in 96-well tissue culture plates at the density of 0.2 × 104 cells per well with 100 μL medium, and were allowed to become attached for 24 h. One hundred microliters of the medium with different

concentrations of Rg5 (e.g., 0μM, 25μM, 50μM, and 100μM) were added to each well. At indicated times, 30 μL MTT stock solution (3 mg/mL) were added to each well. After culturing the cells at 37°C for 2 h, dimethyl sulfoxide (DMSO) was added to dissolve the formazan crystals. MS275 The absorbance was read at the wavelength of 540 nm with a microplate reader (EL800, Biotek Instruments Inc., Winooski, VT, USA). After treatment, the pellet of cells was rinsed with ice-cold phosphate buffered saline (PBS) and lysed in radioimmunoprecipitation assay buffer (0.1% sodium dodecyl sulfate, 0.5% sodium deoxycholate, 50mM Tris-HCl Abiraterone and 0.1% NP-40, pH 8.0 with 150mM sodium chloride) for 1 h at 4°C. The cell lysate was cleared by centrifugation at 17,000 rpm for 10 min at 4°C. Each supernatant sample was separated by 10% sodium dodecyl sulfate–polyacrylamide gel electrophoresis

and the separated protein was transferred to polyvinylidene fluoride (PVDF) membranes. After blocking with 5% nonfat dry milk in TBS-T (25mM Tris and 0.1% Tween 20, 137mM sodium chloride) at room temperature for 2 h, the membranes were incubated with primary antibodies overnight at 4°C and treated with horseradish peroxidase-conjugated secondary antibodies for 2 h. The signals were detected with the ECL Advance Detection Kit (GE Healthcare Bio-Sciences Corp., Piscataway, NJ, USA) by LAS-3000 luminescent image analysis. Apoptosis was evaluated by annexin V/fluorescein isothiocyanate/propidium iodide (annexin V-FITC/PI) dual staining. Treated cells were harvested and resuspended in 1× binding buffer. A combination of annexin V/FITC solution and PI solution were added to each tube. The stained cells were incubated at room temperature for 30 min in the dark. Samples were analyzed by the FACSCanto II Flow Cytometer (BD Biosciences, San Jose, CA, USA).

After antigen uptake, immature DCs become mature and sensitize naive T cells, which leads to clonal expansion and differentiation into effector helper T cells and cytotoxic T cells, which

produce IFN-γ. Mouse DCs treated with ginsenosides in a recent study showed a suppressed maturation process [10]. In mouse DCs stimulated with LPS, the ginsenosides inhibit the secretion of IL-12, an important cytokine that induces T cell activation. However, no reports have revealed Ipatasertib supplier the effect of ginsenosides on the differentiation of immature DCs from human monocytes. In the present study, we therefore explored the effect of ginsenoside fractions on the differentiation of CD14+ monocytes to DCs, and explored the expression of cell surface markers (e.g., CD80, CD86, CD40, and MHC class II) on the differentiated DCs and interferon gamma (IFN-γ) production in CD4+ T cells when cocultured with DCs that were differentiated

in the presence of ginsenoside fractions. Roswell Park Memorial Institute (RPMI) 1640 medium, fetal bovine serum (FBS), and antibiotics (e.g., penicillin and streptomycin) were purchased from Gibco-BRL (Grand Island, NY, USA). Escherichia coli LPS (026:B6), the c-Jun N-terminal kinase (JNK) inhibitor SP600125, and polymyxin B (PMB) were purchased from Sigma–Aldrich (St. Louis, MO, USA). The mitogen-activated protein kinase (MAPK) inhibitor U0126 was purchased from EMD Millipore (San Diego, CA, USA). Human recombinant IL-4, GM-CSF, and anti-Annexin-V-FITC antibody were purchased from R&D Systems (Minneapolis, MN, USA). Rabbit antiphospho-extracellular signal-regulated kinase 1/2 Kinase Inhibitor Library (antiphospho-ERK1/2), anti-ERK1/2, antiphospho-JNK, anti-JNK, antiphospho-p38, anti-p38, and anti-inhibitory kappa B (anti-IκB) antibodies were purchased from Cell Signaling Technology (Danvers, MA, USA). Goat antimouse immunoglobulin G-horseradish peroxidase (IgG-HRP), mouse antirabbit IgG-HRP, and mouse monoclonal anti-β-actin were purchased from Santa Cruz Biotechnology (Santa Cruz, CA, USA).

The specific antibodies for flow cytometric analysis, which included human anti-CD80-PE, anti-CD86-antigen-presenting cell (APC), anti-CD40-fluorescein isothiocyanate (FITC), anti-CD14-FITC, anti-CD11c-APC, and anti-human leukocyte antigen DR (HLA-DR)-FITC were purchased from BD Biosciences (San Diego, PD184352 (CI-1040) CA, USA). Unless otherwise noted, all other reagents were obtained from Sigma-Aldrich (St. Louis, MO, USA). Ginsenoside fractions were extracted from Panax ginseng, as previously described [11]. In brief, the dried root of Panax ginseng was refluxed twice with 80% methanol and concentrated with a vacuum-evaporator. The concentrate was diluted with water and the solution was extracted with 1 L of diethyl ether. The aqueous phase was briefly evaporated under vacuum to remove the remaining ether. The solution was then extracted with n-butanol. The organic phase was finally collected and evaporated.

The result is that the physical attributes of land surface systems more closely reflect unspecified past rather than present conditions,

and that the present state of these systems cannot be easily matched with prevailing climate. In a uniformitarian context, this means that evaluations of system state under present conditions of climatic or environmental forcing cannot be used as a guide to estimate the spatial/temporal patterns or magnitude of past forcing. The logic of this approach is clearly demonstrated in landscapes where cosmogenic dating has been applied to exposed rock surfaces that have been subject to subaerial weathering over long time periods (e.g., Bierman and Caffee, 2001 and Portenga and Bierman, 2011). The dates obtained from this approach span a range of ages showing that, Screening Library ic50 across a single region, land surface weathering does not Ceritinib clinical trial take place at a uniform rate or affect all parts of the landscape equally. The result is a mosaic of landscape palimpsests (Bailey, 2007) in which some landscape elements reflect present-day forcing, whereas others are relict and reflect climatic controls of the past (Stroeven et al., 2002 and Knight and Harrison, 2013b). This shows both the spatial and temporal contingency of geomorphological sensitivity, and that uniformitarian principles

fail to account for the formation of landscape palimpsests, even in the same location and under the same conditions of forcing. Uniformitarianism also

cannot account for the feedbacks associated with system behaviour. For example, over time as ecosystems become established on a sloping land surface, soil thickness increases and hillslope angle decreases due to soil creep. This means that slope systems’ dynamical processes operate at slower rates over time as they converge towards quasi-equilibrium (Phillips, 2009). As a consequence, in this example, system sensitivity to forcing decreases MRIP over time, which is a notion opposed to the steady state and steady rate of change argued through uniformitarianism. Human activity is a major driver of the dynamics of most contemporary Earth systems, and has pushed the behaviour of many such systems beyond the bounds of their natural variability, when based on examination of system dynamics over recent geological time (Rosenzweig et al., 2008 and Rockström et al., 2009). A useful measure of Earth system behaviour is that of sediment yield, which is the product of land surface processes. In many areas of the world, sediment yield has been dramatically increased (by several orders of magnitude above background geological rates) by a combination of human activities including deforestation, agriculture, urbanisation and catchment engineering (Hay, 1994, Wilkinson and McElroy, 2007 and Syvitski and Kettner, 2011).