NI = the

number of intercepts that cross basal membrane,

NI = the

number of intercepts that cross basal membrane, which is proportional to the perimeter of the airway; L = number of points hitting the airway lumen, which is proportional to the intraluminal area. BI was quantified in five non-cartilaginous airways per animal at 400× magnification ( Sakae et al., 1994). Airway smooth muscle area, airway epithelium thickness and edema were defined as the www.selleckchem.com/products/s-gsk1349572.html number of point hitting, respectively, in smooth muscle and epithelial cells and peribronchial edema. This value was divided by the number of intercepts that cross the basal membrane, which is proportional to the perimeter of the airway (Sakae et al., 1994 and Vieira et al., 2007). Measurements were performed in five airways per animal at 1000× magnification. After blood collection from the cava vein, the samples were immediately centrifuged for 15 min (5 °C; 1000 rpm). Serum samples were stored at −70 °C until the DAPT clinical trial assay was performed. A PCA reaction was used to detect and estimate the levels of anaphylactic IgE and IgG1 OVA-specific antibodies as previously described (Ovary, 1964 and Mota and Perini, 1970). Briefly, the back of a naïve guinea

pig was shaved, and 0.1 ml of different serum dilutions was injected intradermally. Thirty naïve guinea pigs were used to evaluate the PCA, and the serum from each animal was included in the study (n = 30). After a long latent period of 48 h for IgE or a short period of 24 h for IgG1, the animals were challenged intravenously (i.v.) with 1 ml of a 0.5% solution of Evans blue in saline (0.9% NaCl) containing 1 mg of antigen (ovalbumin). The animals were

euthanized 30 min after injection of the antigen, and the diameters of the blue spots on the inner surface of the flayed skin were measured. To detect the IgG1-type antibody, the serum was heated for 3 h at 56 °C to inactivate IgE activity; the heated serum was injected for PCA after a short latency period. The PCA titers were defined as the highest dilutions that gave an intradermal allergic Resveratrol reaction larger than 5 mm in diameter in triplicate tests ( Ovary, 1964 and Mota and Perini, 1970). One-way analysis of variance (ANOVA) followed by a Student–Newman–Keuls post hoc test (parametric data) or ANOVA on ranks followed by Dunn’s post hoc test (non-parametric data) were used to compare the different parameters between groups. The values were expressed as the mean ± SD for parametric data and as the median (variance) for non-parametric data. The level of significance was set at p < 0.05. Table 1 shows the maximal exercise capacity obtained in initial and final tests for each group before and after the AE protocol. Only animals from the trained groups (AE and OVA + AE groups) exhibited a significant increase in exercise capacity when compared with the animals in the non-trained groups (C and OVA) (p < 0.001; Table 1). The OVA and OVA + AE groups had increases in the OVA-specific IgE and IgG1 titers compared to the non-sensitized groups (p < 0.001; Table 1).

, 2008, Rick et al , 2009b and Rick, 2013) Fig 2a documents the

, 2008, Rick et al., 2009b and Rick, 2013). Fig. 2a documents the BTK inhibitor screening library timing of some human ecological events on the Channel Islands relative to human population densities. We can say with confidence that Native Americans

moved island foxes between the northern and southern Channel Islands ( Collins, 1991 and Vellanoweth, 1998) and there is growing evidence that humans initially introduced mainland gray foxes to the northern islands ( Rick et al., 2009b). Genetic, stable isotope, and other studies are under way to test this hypothesis. Another island mammal, Peromyscus maniculatus, appears in the record on the northern Channel Islands about 10,000 years ago, some three millennia after initial human occupation, and was a likely stowaway in human canoes ( Walker, 1980, Wenner and Johnson, 1980 and Rick, 2013). On the northern Channel Islands, Peromyscus nesodytes, a larger deer mouse had colonized the AZD6244 chemical structure islands prior to human arrival, sometime during the Late Pleistocene. The two species of mice co-existed for millennia until the Late Holocene when P. nesodytes went extinct, perhaps related to interspecific competition with P. manicualtus and changing island habitats

( Ainis and Vellanoweth, 2012 and Rick et al., 2012a). Although extinction or local extirpation of island mammals and birds is a trend on the Channel Islands, these declines appear to be less frequent and dramatic MYO10 than those documented on Pacific and Caribbean Islands, a pattern perhaps related to the absence of agriculture on the Channel Islands and lower levels of landscape clearance and burning (Rick et al., 2012a). Fires have been documented on the Channel Islands during the Late Pleistocene and Holocene (Anderson et al., 2010b and Rick et al., 2012b), but we are just beginning to gain an understanding of burning by the Island Chumash. Ethnographic sources document burning by Chumash peoples on the mainland (Timbrook et al., 1982), but say little about the islands. Anderson et al. (2010b) recently presented a Holocene record

of fire history on Santa Rosa Island, which suggests a dramatic increase in burning during the Late Holocene (∼3000 years ago), attributed to Native American fires. Future research should help document ancient human burning practices and their influence on island ecosystems. For now, we can say that the Island Chumash strongly influenced Channel Island marine and terrestrial ecosystems for millennia. The magnitude of these impacts is considerably less dramatic than those of the ensuing Euroamerican ranching period (Erlandson et al., 2009), a topic we return to in the final section. Archeological and paleoecological records from islands provide context and background for evaluating the Anthropocene concept, determining when this proposed geological epoch may have begun, and supplying lessons for modern environmental management.

Dedicated lures for this purpose have been developed ( Mands et a

Dedicated lures for this purpose have been developed ( Mands et al., 2004) and commercial traps have been produced, although the efficacy see more of these in reducing overall Culicoides biting rates (and hence arbovirus transmission risk) has not been quantified in Europe. To date, lures have not been developed for livestock-associated Culicoides species, although preliminary studies have been conducted with generic attractant compounds that show promise ( Harrup et al., 2012). In the case of C. impunctatus, these techniques are unlikely

to lead to permanent reductions in population abundance due to autogeny and huge source populations, but they may impact on the major species associated with livestock, which are largely anautogenous. In the event of an incursion of an arbovirus into Europe that is capable of person-to-person spread by Culicoides midges, education is likely to play a key role in promoting avoidance of areas supporting Metformin substantial populations of vectors. The substantial nuisance already inflicted by C. impunctatus has led to the development of a ‘midge forecast’ for tourists and local inhabitants in Scotland which is disseminated

via newspapers, a website (http://www.midgeforecast.co.uk/) and most recently a mobile phone application. Combined with data concerning C. impunctatus distribution and fine-scale habitat use, the midge forecast could be usefully employed to warn of geographical areas and habitats of high exposure risk. A clearer understanding of Protein tyrosine phosphatase which recreational activities and jobs require prolonged exposure to Culicoides would be extremely useful in qualitatively assessing risk of exposure. Overlap on farms between Culicoides populations and human workers is more difficult to assess, however, and may be significantly influenced by husbandry practices. For example, it is quite possible that those involved in forestry or game-related activities in proximity to farms may suffer greater exposure than the farmers

themselves. Culicoides are among the most abundant vectors of arboviruses found in Europe, but current evidence demonstrates that their impact on human health in this region is currently limited to biting nuisance. However, the existence of one or more proven, but as yet undescribed, route of entry for Culicoides-borne arboviruses into Europe raises the potential of future impacts on human health. From reviewing current knowledge of Culicoides populations both in Europe and in areas of arbovirus transmission worldwide we reach the following conclusions: 1. Future introduction of known or unknown arboviruses that are transmitted in epidemics between humans by Culicoides (OROV) are unlikely to lead to sustained outbreaks of disease in Europe without the involvement of additional vector groups and/or as yet unknown reservoir hosts.

Cousin et al [14] reported that the level of lipids in the perito

Cousin et al [14] reported that the level of lipids in the peritoneum increased after denervation. This suggests that the sympathetic nervous system influences the activity or differentiation

CHIR-99021 solubility dmso of white adipose cells. Parasympathetic nerves as well as sympathetic nerves showed a relation with adipose tissue. Kreier et al [15] reported that when the parasympathetic nerve was removed, the HSL activity in white adipose tissue increased. However, the absorption of FFA and blood glucose decreased. Given that the direct measurement of autonomic nervous activity by micrography is not feasible in a large epidemiological study, heart rate variability (HRV) is used as the measurement method for the autonomic nervous system [16]. HRV is measured by the variation of the beat-to-beat interval. The average R-R is calculated

by 60 divided by the average heart rate in beats/min. Chang et al [17] showed that HRV is related to several component of metabolic syndrome (MtS). When they separated 1,298 individuals into four groups based on the components ABT 888 of MtS, those who had one or more components of MtS showed a lower standard deviation of the R-R interval compared to a healthy control group. The recorded use of ginseng dates back 2,000 years. It has been one of the most popular herbal supplements in Asia, especially in Korea, China, and Japan. In the USA, ginseng ranked as one of the top-10 selling herbal supplements in 2003 [18]. The primary effective components of ginseng are known as ginsenosides, and these include compound K (CK), Rg3, Rk1, and Rg5, all of which have a steroidal skeleton. In the results of this study,

CK served as the ligand of glucocorticoid receptor (GR) [19] and Rg3 functioned as the ligand of estrogen receptor (ER) [20], which implies a possible effect of ginseng on lipolysis. In fact, when CK was administered to a 3T3-L1 adipocyte cell line, the storage of triglycerides was suppressed. On the other By contrast, Rg1 stimulated triglyceride storage in adipocytes [21]. Rg3, Rk1, and Rg5 treatments in oxyclozanide 3T3-L1 suppressed lipid accumulation [22]. As well as the reported effects of ginseng on FFA, red ginseng has also been shown to have a beneficial effect on insulin and glucose regulation. Vuksan et al [23] reported that red ginseng consumption improved insulin and glucose regulation in type 2 diabetes patients. Lee et al [24] showed that red ginseng has a beneficial effect on insulin sensitivity. We also reported that fermented red ginseng (FRG) showed a serial causal effect on the level of hormones, insulin resistance, and insulin levels. In an analysis of the effects of hormones on glucose blood levels, the difference between the FRG group and the placebo group was due to the level of aldosterone [25]. According to an experiment with mice, ginsenosides stimulated an acetylcholine release in the terminal of cholinergic neurons [26].

10 Despite the amount of uncertainty placed on these volume–weig

10. Despite the amount of uncertainty placed on these volume–weight calculations it appears

see more that published C-factors nonetheless underestimate the effects of urban forest cover in the region ( Fig. 11); however, in order to elucidate an appropriate C-value range for the area, an assessment of the contributions to the pond’s sediment budget unaffiliated with sheet and rill erosion from the surrounding landscape is required. The following two sources offer explanation for the inclusion of materials not accounted for by the USLE, which contribute to the overestimation of pond sedimentation due to inter-rill and rill processes: (1) sediment transported into the pond by anthropogenic means, and (2) gully erosion from surrounding hillsides. The pond is the final resting place for agonist all materials derived from surrounding hillslopes and the footpath. A small source of error that could explain some of the variance between field and numerical model results is presented by the unknown factors associated with the upkeep of the footpath around the pond. Sand and gravel are replaced here on a regular basis as hillslope runoff not only carries materials from the slopes,

but also from the footpath into the pond. Evidence for this process is found in cores, which contained scattered pebbles found concentrated on the footpath. Since no records exist that would allow for quantification of this sediment source, the extent to which Metalloexopeptidase these materials

offset measurements cannot be pursued; however, based on an assessment of collected sediment cores and a comparison of pond-sediment volume against path dimensions, it is assumed this contribution is negligible. It is likely that gullies are a significant contributor to the USLE model deviation; however, they provide an unquantified volume of sediment to the pond’s budget and pursuing their contribution from a process-oriented perspective would be time-consuming. It is estimated based on field reconnaissance of gully dimensions (width and depth) and their extent (derived from the flow-accumulation model) that the volume represented by gullies along the steep slopes north of the pond corresponds to ∼100 m3. This Gully-volume estimate is an order of magnitude smaller than the volume of sediment emplaced into the pond (∼6228 m3) and therefore would do little to close the gap between USLE model estimates of inter-rill and rill erosion and quantified pond sedimentation (Fig. 11). Regardless of how much gullying and anthropogenic contributions may add to the pond’s sediment budget, evidence suggests that urban forest cover promotes high erosion rates, which translate to high sediment flux and deposition within the pond. This is a function of the absent sediment storage anywhere along route within the watershed (Fig. 3); the study area thus provides a suitable location for a qualitative assessment of the C-value for this land-cover type.

Humans hunted seals and sea lions since at least the Terminal Ple

Humans hunted seals and sea lions since at least the Terminal Pleistocene, but early records of pinniped hunting are scarce, with dramatic increases at some locations beginning around 1500 years ago ( Braje et al., 2011a, Braje et al., 2011b and Erlandson et al., 2013). One of the more interesting trends in

pinniped demographics during the Holocene compared to today is the changing abundance of Guadalupe fur seals and elephant seals ( Fig. 2c; Rick et al., 2009a and Rick et al., 2011). For much of the Holocene, Guadalupe fur seals are the most abundant taxa found in archeological sites, suggesting they were frequently encountered when hunting and scavenging. In contrast, elephant seals are rarely found in archeological sites, with just a handful of bones found in island (or mainland) sites. Both of these species were hunted to near Selleckchem Sunitinib extinction during the 18th–19th century global fur and oil trade. Following federal protection in the 1970s, populations have grown exponentially and

there are now more than 50,000 elephant seals in Alta California waters. Guadalupe fur seals, however, are very rare north of Nutlin-3 order Mexico, with only a few observations during the last decade ( Rick et al., 2009a). These dramatic differences in abundance between Holocene seal and sea lion populations and those of today suggest that recovered pinniped populations are not ‘natural’ and are largely an artifact of management and conservation (see Braje et al., 2011a, Braje et al., 2011b and Erlandson et al., 2013). Seal and sea lion conservation can lead to debate between conservationists focused on the management of marine mammal populations and commercial fisheries concerned about shellfish and fish stocks that are common prey of pinnipeds and sea otters. Such conflicts have also begun in Hawaii with debate over monk seal conservation and the effects on Hawaiian fisheries and recreation. Finally, the extensive growth of some pinniped

populations in California demonstrates the conflicts between natural and cultural resource management, with pinnipeds hauling filipin out on, disturbing, and destroying non-renewable archeological sites located on the shoreline of the Channel Islands and elsewhere (see Braje et al., 2011a and Braje et al., 2011b). The records of finfish and seabirds are just beginning to be explored in detail, but Braje et al. (2012) recently documented size changes in rockfish (Sebastes spp.), including estimates that many prehistoric specimens were larger than modern fishes. Chendytes lawi, an extinct flightless duck, appears to have been slowly pushed to extinction on the Channel Islands and mainland by human predation and other variables over several millennia ( Jones et al., 2008 and Rick et al., 2012a). Along with human hunting, the extinction of C.

, 2007 and Steffen et al , 2011) suggested that AD 1800, roughly

, 2007 and Steffen et al., 2011) suggested that AD 1800, roughly the start of the Industrial Revolution in Europe, be considered as the beginning of the Anthropocene. Others have taken a longer view, especially Ruddiman, 2003, Rigosertib nmr Ruddiman, 2005 and Ruddiman, 2013, who argued that greenhouse gas concentrations, deforestation, soil erosion, plant and animal extinctions, and associated climate changes all accelerated at least 8000 years ago with wide-scale global farming (see also Smith and Zeder, 2014). Doughtry et al. (2010) suggested that the Anthropocene should be pushed back to 14,000 or 15,000

years ago, eliminating the Holocene, and correlating with the extinction of Pleistocene megafauna and the associated climate changes brought on by these events. At the other end of the spectrum, some scholars argue for a starting date of AD 1950, based on changes in riverine fluxes (Maybeck and Vörösmarty, 2005) or the appearance of artificial radionucliotides resulting from atomic detonations (Crutzen and Steffen, 2003). In 2008, a proposal

for the formal designation of the Anthropocene was presented to the Stratigraphy Commission of the Geological Society of London (Zalasiewicz et al., 2008). An Anthropocene Working Group, part of the Subcommission on Quaternary Stratigraphy, has been formed to Tanespimycin solubility dmso help determine if the Anthropocene will be formally accepted into the Geological Time Scale and when it began (Zalasiewicz et al., 2010,

p. 2228). In line with Crutzen’s arguments, the proposal suggests a genesis at the dawn of the Industrial Revolution or the nuclear era of the 1950s. Ultimately, any date chosen for the beginning of the Anthropocene is likely to be relatively arbitrary and controversial, a point at which scientists can logically argue that we have moved from a planet dominated by natural processes into one dominated by anthropogenic forces. No single date can do justice, moreover, to the long process of human geographic expansion, technological Epothilone B (EPO906, Patupilone) development, and economic change that led up to the Industrial Revolution, the nuclear age, or any other singular hallmark in planetary history. As demonstrated by the papers in this issue, archeology—the study of material remains left behind by past human cultures—has much to contribute to understanding the deep history of human impacts on earth’s landscapes and ecosystems. From the controversial and often polarized debates about the history of anthropogenically driven extinctions, to the origins and spread of agricultural and pastoral societies, the effects of humans on marine fisheries and coastal ecosystems, to the acceleration of colonialism and globalization, archeological records can be utilized by scholars to understand not just when humans dominated earth’s ecosystems, but the processes that led to such domination.

2F–J) Most of the proton-generating processes are associated wit

2F–J). Most of the proton-generating processes are associated with the cultivation-induced changes in organic-matter cycles, typically the loss of organic matter from the soil owing to the increased Fluorouracil purchase organic-matter decomposition and product removal. In this study, the ginseng planting obviously reduced the TOC concentrations of ginseng soils, which is positively correlated with the pH (r = 0.293, p < 0.05, n = 60). The decrease in the TOC is one of the causes of the decreased pH. Base cations were investigated seasonally (Fig. 1A–T). Ginseng planting had negligible effects on the concentrations of Ex-Na+, Ex-K+, and exchangeable Mg2+. The elevated concentrations

of Ex-Na+ and Ex-K+ in the next spring

may have been derived from the release of exchangeable metal ions bound to strong cation exchange sites on the surface of soil minerals left by frost. There was, however, a remarkable decrease in the concentration of Ex-Ca2+ (Fig. 1A–T). Considering the vegetation age and temporal variation, we propose that ginseng might require more Ca to grow. Konsler and Shelton [10] found that ginseng plants took up Ca MG-132 datasheet more readily in soils. Ca deficiencies can be seen in stunted ginseng that lack general vigor and have smaller and more fragile growth buds [21]. Soil Ca has also been proposed as a key element in the success of American ginseng crops in forest soils [22]. Wild populations of American ginseng in the United States are found in a wide range of soil pHs but always in Ca-rich soils [23]. Beyfuss even found that healthy populations of wild ginseng grew in soil conditions with very low pH and very high levels of Ca [24], which is abnormal in mineral soils. In this study, the decrease in Ex-Ca2+ in the bed soils added new evidence that Asian ginseng needs more Ca to grow and that Ca is the key factor for successfully planting Asian ginseng. Furthermore, the Ex-Ca2+ concentrations positively correlated with the pH (r = 0.325, p < 0.01, n = 60)

within the ginseng bed. The decrease in Ex-Ca2+ concentrations might be one of the factors resulting in pH decreases in bed soils ( Fig. 1 and Fig. 3A–E). It is well known that the soil pH has a large Rebamipide influence on ginseng growth and development [10] and [11]. Red skin indices of ginseng were reported to agree well with the Al3++H+, Al3+ levels [11]. In acidic soils, most plants become stressed as result of a toxic concentration of Al3+[25]. Both low Ca and high Al concentrations were measured in the soils of American ginseng fields, and Ca deficiency and Al toxicity were proposed to have resulted in the higher susceptibility of American ginseng to abiotic and biotic stresses [22]. A risk assessment for Al toxicity in forests has also been based on different methods using soil- and/or plant-based indices [26].

In addition to their numerical dominance, many oligarchs have the

In addition to their numerical dominance, many oligarchs have the ability to persist under intensive, long-term exploitation without depletion. Their communities tend to be rich in younger individuals, reflective of their successional nature and reproductive ability. Even compared to industrial agriculture, the oligarchic species produce a tremendous amount of food, materials, and revenue (Peters et al., 1989). For example, the prehistoric Brazil-nut-dominated groves were the basis for most of Brazil’s export sales for decades (Smith et al., 1992:384–402), and well-managed anthropic acai

groves yield significant income for long periods (Brondizio, 2009 and Goulding and Smith, 2007:121–146), in addition to AT13387 clinical trial an important food supplement. Moriche, also, is a long-lived palm highly productive of fruit and trunk starch or sap (Fig. 14) (Goulding and Smith, 2007:51–120; Peters et al., 1989). Because so few archeological sites of the Amazon cultural sequence have been sampled intensively for botanical remains, we know little about the human history and development of forests in different regions. Paleoindian botanical remains and stable carbon isotopes suggest that those first colonists initiated the development of cultural forests in Amazonia, in the form of upland palm forests (Section ‘Environmental impacts of early nomadic foragers’). We also know that forest structure changed distinctly through time in the catchments of later settlements

(Roosevelt, 2000:472–476,

480–486). On Marajo, stable carbon isotopes of human bone and carbonized plants are more than five per mil less negative during late prehistory, see more suggesting that locally forests became more open around the long-term mound villages. At Santarem, though Formative people had access to slow-growing, closed canopy forests for food and fuel, people of the large late-prehistoric black soil site there used more open and fast-growing forest. The stable carbon isotope ratios show a several per-mil change from the Formative woods to those of late prehistory, consistent with thinning of forest canopy. Preliminary identifications of the carbonized Methamphetamine wood include fast-growing tree species of the Rutaceae family common in agroforestry plots and secondary forest around Santarem today (Roosevelt, 2000). Achieving a more accurate map of the cultural forests is an important goal. They are potential evidence of the Anthropocene but also a significant economic resource of food and raw materials and a repository of natural and cultural diversity (Anderson and Posey, 1989, Clement, 1999, Goulding and Smith, 2007, Peters et al., 1989 and Smith et al., 2007). The oligarchic forests can produce for hundreds of years, yielding fruits and nuts for subsistence, for both local and global markets, wood for fuel and construction, materials for tools, fabrics, and containers, and vegetation cover needed to ameliorate the temperature and moisture extremes of the tropical climate.

5 m below m s l This area became a lagoon much later than the mo

5 m below m.s.l. This area became a lagoon much later than the more northern and southern parts, where the sea arrived about 7000 BP ( Canali et al., 2007) and about 6000 cal years BP ( Zecchin et al., 2009), respectively. In correspondence

with reflector (2), the salt marsh facies Lsm reveals the presence of a buried salt marsh (alternatively emerged and SB203580 submerged) overlaid by the mudflat facies Lm (in green in Fig. 2a). At 2.21 m, 1.89 m and 1.5 m below m.s.l., three calibrated 14C ages (Table 1) of peat and vegetal remains samples collected in salt marsh, intertidal and subtidal environments, respectively allowed us to reconstruct the evolution of the salt marsh. There was a salt marsh during the Iron Age going back to 863 BC that still existed in 459 BC (before the first stable settlements in the lagoon islands), being sometimes submerged. The salt marsh had disappeared by 240 AD during Roman Times. Core SG24 intersects a large palaeochannel (CL1, Fig. 2 and Fig. 3). The reflection pattern of the palaeochannel is about 110 m wide and extends vertically from about 2 m to about 6 m under the

bottom. The lowest high-amplitude oblique reflector corresponds to the transition from the laminated channel facies Lcl and the sandy channel facies Lcs that is not penetrated by the high frequency acoustic signal as already observed in Madricardo et al. (2007). The channel infill structure includes oblique clinoforms that are sub-parallel and of high-to-moderate amplitude. They have moderate-to-low continuity, dipping southward in the northern part of the palaeochannel. They correspond to the difference of Ipatasertib in vitro acoustic impedance between layers of clayey silt and thin sandy layers within the tidal channel facies Lcl. This configuration is the result of the active lateral accretion through point bar migration of a large meander palaeochannel in an area that is now a submerged mudflat. The angle of the clinoforms decreases southwards suggesting

a phase of lower energy and decreased sediment grain-size. A slightly wavy low amplitude horizon at about 3 m below m.s.l. suggests the decrease or even the end of the activity of the channel. The 14C dating of plant remains at 6.56 m below m.s.l. in a highly energetic channel environment indicates Amobarbital that the channel was already active at 819 BC. Therefore, the channel was active at the same time as the salt marsh before the first human settlements in the lagoon. The 14C dating of a shell at 2.61 m below m.s.l. in a subtidal environment confirms that the channel ceased activity in this site by 365 BC. In the upper part of the profile (for about 2 m beneath the bottom) the acoustic pattern is chaotic. This chaotic upper part corresponds to the sedimentary facies of the mudflat Lm in core SG24 (in green in Fig. 2). The study of the acoustic and sedimentary facies of the palaeochannel CL2 (in profile 2, 3 and 4 and cores SG25, SG27 and SG28 in Fig.