Here we propose a dimensionless metric to help identify when a ch

Here we propose a dimensionless metric to help identify when a channel is incised, “relative incision,” that quantifies ht/de, the ratio of terrace height (ht) relative to effective flow depth (de). Field data show that average bar height in Robinson Creek is 0.6 m; thus, effective flow depth is inferred to

be 0.85 m above Bleomycin supplier the thalweg. In Robinson Creek the relative incision ratio ranges from 8.0 to 13.3 in the upstream and downstream portion of the incised study reach, respectively. In contrast, in a stable alluvial channel without incision, the floodplain height would approximate the depth of the effective discharge necessary to transport bed material and form bars and the relative incision ratio would be 1.0. Thus, as a channel incises, a gradient of diminishing connectivity

and increased transport capacity accompanies an increase in relative incision above a value of 1.0. Quantifying the metric is useful because identifying alluvial incision implies that we can unambiguously differentiate an incised channel from a non-incised channel. In particular, other fluvial characteristics, such as eroding vertical stream banks, sometimes make identification via visual observation difficult within naturally highly variable and to varying degrees disturbed “Anthropocene” fluvial systems. Further work is warranted to distinguish floodplain from terrace landforms to assess the importance of incision as a formative geomorphic process, especially when relative incision ratios are close to

agonist 1.0. The magnitudes and rates of channel incision characteristic of the “Anthropocene” are unprecedented in geologic time in the absence of driving mechanisms such as climate change that modifies a watershed’s hydrology and sediment supply, sea level lowering that changes baselevel, or tectonic events that modify Methane monooxygenase channel slopes. As an illustration of the problem, the field study of Robinson Creek in Mendocino County, California, suggests spatially diverse causes of incision. They include land use changes such as grazing beginning in about 1860 that likely changed hydrology and sediment supply, downstream baselevel lowering over the same temporal period, and local channel structures built to limit bank erosion. Channel incision in Robinson Creek likely progressed during episodic floods that recur on average during 25% of years. Bank heights average 4.8–8.0 m, from the upstream to downstream end of a 1.3 km study reach. Development of the “relative incision” ratio of terrace height (ht) to effective flow depth (de) as a metric to quantify incision yields values of 8.0–13.3 times the threshold value of 1.0. Further work is warranted to compare magnitude of incision in Robinson Creek other incised or stable systems. Incision leads to significant ecological effects such as destabilization of riparian trees and loss of channel-floodplain hydrologic connectivity.

, 2005a, Erlandson et al , 2005b and Rick et al , 2008a) By 7000

, 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) find more 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 Antidiabetic Compound Library 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 Thymidylate synthase 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.

New competitors and predators were introduced from one end of the

New competitors and predators were introduced from one end of the globe to the other, including rodents, weeds, dogs, domesticated plants and animals, and everything in between (Redman, 1999:62). Waves of extinction mirrored increases in human population growth and the transformation

of settlement and subsistence systems. By the 15th and 16th centuries AD, colonialism, the creation of a global market economy, and human translocation of biota around the world had a homogenizing effect on many terrestrial ecosystems, disrupting both natural and cultural systems (Lightfoot et al., 2013 and Vitousek et al., 1997b). Quantifying the number and rates of extinctions over the past 10,000 years is challenging, however, as global extinction rates are difficult to determine even today, in part because the majority of earth’s species still remain undocumented. GDC-0941 purchase The wave of catastrophic plant and animal extinctions that began with the late Quaternary megafauna of Australia, Europe, and the Americas has continued Bortezomib mouse to accelerate since the industrial revolution. Ceballos et al. (2010) estimated that human-induced species extinctions are now thousands of times greater than the background extinction rate. Diamond (1984) estimated that 4200 (63%)

species of mammals and 8500 species of birds have become extinct since AD 1600. Wilson (2002) predicted that, if current rates continue, half of earth’s plant and animal life will be extinct by AD 2100. Today, although anthropogenic climate change is playing a growing role, the primary drivers of modern extinctions appear to be habitat loss, human predation, and introduced species (Briggs, 2011:485). These same drivers contributed to ancient megafaunal and island extinctions – with natural forces gradually giving way to anthropogenic changes – and accelerated after the spread of domestication, agriculture, urbanization, and globalization. In our view, the acceleration

of plant and animal extinctions that swept the globe beginning after about 50,000 years ago is part of a long process that involves climate change, the reorganization of terrestrial ecosystems, human hunting and habitat alteration, and, Carteolol HCl perhaps, an extraterrestrial impact near the end of the Pleistocene (see Firestone et al., 2007 and Kennett et al., 2009). Whatever the causes, there is little question that the extinctions and translocations of flora and fauna will be easily visible to future scholars who study archeological and paleoecological records worldwide. If this sixth mass extinction event is used, in part, to identify the onset of the Anthropocene, an arbitrary or “fuzzy” date will ultimately need to be chosen. From our perspective, the defined date is less important than understanding that the mass extinction we are currently experiencing has unfolded over many millennia.

We found that the ability to represent recursion in the visual do

We found that the ability to represent recursion in the visual domain was selleck chemical correlated with grammar comprehension, and that this correlation was partially independent from general intelligence. However this effect was not specific to recursion, since grammar comprehension also correlated with embedded iteration. This suggests that grammar comprehension abilities were correlated with a more general ability to represent and process hierarchical structures generated

iteratively, independently of whether these were recursive or not. This result is not completely surprising given that not all syntactic structures in TROG-D are recursive, although all are hierarchical. We also assessed whether there was a more specific correlation between visual recursion and embedded clauses, but found again only a general association with both EIT and VRT. However, it is important to note that TROG-D only includes sentences with one level of embedding, e.g. relative clause (nominative): Der Junge, derdas Pferd jagt, ist dick ‘The boy, who is chasing the horse, is chubby’. Children may potentially use non-recursive representations for these kind of sentences ( Roeper, 2011). Only a task focussed on sentences with several levels of recursive embedding would allow a direct comparison between visual

recursion and syntactic recursion. Despite this limitation, it is interesting that performance on our novel AZD2281 price visual tasks was correlated with grammar abilities, even when the effects of non-verbal intelligence were taken into account. These correlations could be explained by the existence of shared cognitive resources, independent from non-verbal intelligence, used for the processing of hierarchical structures in both language and visuo-spatial reasoning, or even by the effects of literacy ROS1 (which are partially independent of intelligence) in the processing of hierarchical structures. Interestingly, while individual differences in intelligence predicted VRT and EIT scores both between and within grades, grammatical

comprehension abilities accounted only for differences between grades. Again, this argues in favor of a general age-related maturational influencing the processing of hierarchical structures, occurring between second and fourth grade, which is partially independent from non-verbal intelligence. Furthermore, in our sample, grammar comprehension and non-verbal intelligence were not significantly correlated. Hence, this general maturation process in hierarchical processing cannot be explained solely by the increase of intelligence with age. Future studies with a more comprehensive assessment of grammar (that includes recursion at several levels), and the inclusion of more cognitive tests (assessing cognitive control, attention, etc.) in the experimental procedure could potentially shed more light on a possible relationship between grammar and processing of complex visual structures.

Recognition of the tremendous contributions

of anthropoge

Recognition of the tremendous contributions

of anthropogenic sediment to modern sediment budgets by early geomorphologists (Gilbert, 1917, Happ et al., 1940 and Knox, 1972) led to a fundamental reconsideration of sediment sources in many fluvial environments. Theories of sediment delivery and storage that blossomed in the 1970s, coupled with the recognition of massive loadings of anthropic sediment, DAPT lead to the inescapable conclusion that many fluvial systems are highly dynamic and not in equilibrium with regards to a balance between sediment loads and transport capacity (Trimble, 1977). For example, high sediment loadings in streams of the Atlantic Coastal Plain of the northeastern USA are better explained by recruitment of anthropogenic sediment from floodplains and terraces than by intensive upland land use (Walter and Merritts, 2008 and Wohl and Rathburn, 2013). The awareness of anthropogenic sediment has a long history, although the deposits have been referred to by various names. In many regions of North America, sedimentary deposits were produced by accelerated erosion associated with intensive land clearance

and agriculture following EuroAmerican settlement (Happ et al., 1940, Happ, 1945, Knox, 1972, Knox, selleck chemical 1977, Knox, 1987, Knox, 2006, Trimble, 1974, Costa, 1975, Magilligan, 1985, Jacobson and Coleman, 1986, Faulkner, 1998, Lecce and Pavlowsky, 2001, Florsheim and Mount, 2003, Jackson et al., 2005, Walter and Merritts, 2008, Gellis et Thiamine-diphosphate kinase al., 2009, Merritts et al., 2011 and Hupp et al., 2013). Mining also generated large sedimentation events in North America (Gilbert, 1917, Knox, 1987, James, 1989, Leigh, 1994, Lecce, 1997, Stoughton and Marcus, 2000, Marcus et al., 2001, Bain and Brush, 2005 and Lecce et al., 2008). These anthropogenic deposits are being increasingly referred to as ‘legacy sediment’ (LS) by environmental scientists. Anthropogenic sediment does not

occur uniformly over the landscape but collects in certain locations where it creates landforms. Types of LS deposits vary greatly from colluvial drapes on hill sides, to aprons and fans at the base of hill slopes, to a variety of alluvial depositional features in channels, floodplains, deltas, lakes, and estuaries. (‘Colluvium’ is used broadly in this paper to include mass wasting as well as sheetflow and rill deposits on or at the base of hillslopes (Fairbridge, 1968). It does not necessarily connote anthropogenically produced sediment (LS) as may be implied in central Europe (Leopold and Völkel, 2007).) A typology of LS is described based on locations and geomorphology of deposits. Explanations for heterogeneous spatial patterns of LS deposits are given based on differences in sediment production, transport capacity, accommodation space in valley bottoms, and other factors that are intrinsically geomorphic.

Londoño (2008) highlighted the effect of abandonment on the Inca

Londoño (2008) highlighted the effect of abandonment on the Inca agricultural terraces since ∼1532 A.D., represented by the development of rills and channels on terraces where the vegetation is absent. Lesschen et al. (2008) underlined the fact that that terracing, although intended as a conservation practice, enhances erosion (gully erosion through the terrace walls), especially after abandonment. These authors carried out a study in the Carcavo basin, a semi-arid area in southeastern Spain. More

than half of the abandoned fields in the catchment area are subject to moderate and severe erosion. According to these studies, the land abandonment, the steeper terrace slope, the loam texture of the soils, the valley bottom position, and the presence of shrubs on the terrace walls are all factors that increase the risk of terrace failure. Construction of new terraces should therefore be carefully planned see more and be built according to sustainable design criteria (Lesschen et al., 2008). Lesschen et al. (2008) provided guidelines to avoid the land erosion due to abandonment. They suggested the maintenance of terrace walls in combination with an increase in vegetation cover on the terrace, and the re-vegetation of indigenous grass species on zones with concentrated flow to prevent gully erosion. Lesschen et al. (2009) simulated the runoff

and sediment yield of a landscape scenario without agricultural terraces. They found values higher by why factors of four and nine, respectively, when compared to areas with terraces. Meerkerk et al. (2009) examined SB431542 supplier the effect of terrace removal and failure on hydrological connectivity and peak discharge in a study area of 475 ha in southeastern Spain. They considered three scenarios: 1956 (with terraces), 2006 (with abandoned terraces), and S2 (without terraces). The analysis

was carried out with a storm return interval of 8.2 years. The results show that the decrease in intact terraces is related to a significant increase in connectivity and discharge. Conversely, catchments with terraces have a lower connectivity, contributing area of concentrated flow, and peak discharge. Bellin et al. (2009) presented a case study from southeastern Spain on the abandonment of soil and water conservation structures in Mediterranean ecosystems. Extensive and increasing mechanization of rainfed agriculture in marginal areas has led to a change in cropping systems. They observed that step terraces have decreased significantly during the last 40 years. Many terraces have not been maintained, and flow traces indicate that they no longer retain water. Furthermore, the distance between the step terraces has increased over time, making them vulnerable to erosion. Petanidou et al. (2008) presented a case study of the abandonment of cultivation terraces on Nisyros Island (Greece).

Specifically, we examined choices in the Opt-Out task, in which p

Specifically, we examined choices in the Opt-Out task, in which participants could make a nonbinding choice for LL but could choose SS at any point during the delay period. Since the SS was also available during the initial choice (Figure 1D), and at the time of choice participants knew the delay length, choices for SS during the delay period are suboptimal in terms of maximizing reward across time. Figure 2B displays the proportion of SS choices during

the delay period conditional on initial choices for LL. We observed a substantial number of preference reversals (one-sample t test, Study 1: t(57) = 4.99, p < 0.0001; Study 2: t(19) = 3.94, p = 0.001), which increased as a function of delay (Study 1: F(2,82) = Saracatinib chemical structure 12.50, p < 0.0001; Study 2: F(2,32) = 9.64, p = 0.001; Figure 2B). Preference reversals were positively correlated with the proportion of MAPK inhibitor SS choices in the willpower task at a trend level in Study 1 and significantly so in Study

2 (Study 1: r = 0.251, p = 0.068; Study 2: r = 0.648, p = 0.002). Despite the fact that all tasks had equivalent rewards and delays, self-control differed across tasks (Study 1: F(3,171) = 17.51, p < 0.001; Study 2: F(3,60) = 7.209, p < 0.001; Figure 2C). The opportunity to precommit improved self-control: participants were more likely to choose LL in the Precommitment task than in the Opt-Out task (Study 1: Tau-protein kinase t(57) = 5.64, p < 0.001; Study 2: t(19) = 3.45, p = 0.003) and the Willpower task (Study 1: t(57) = 5.26, p < 0.001; Study 2: t(19) = 3.58, p = 0.002), as well as the Choice task in Study 1 (Study 1: t(57) = 3.40, p = 0.001). Although the mean proportion of LL choices in the Precommitment task was greater than in the Choice task in Study 2, the difference was not significant (t(19) = 1.00, p = 0.328), likely due to the

reduced sample size compared with Study 1. The task-related pattern of choices was consistent across delays (i.e., the task × delay interaction was not significant, Study 1: F(6,342) = 1.16, p = 0.330; Study 2: F(6,114) = 1.10, p = 0.369). The improvement in self-control observed in the Precommitment task varied across subjects, such that more impulsive individuals were more likely to benefit from precommitment. We defined impulsivity, here, as breakdown of willpower; impulsivity was therefore estimated as the proportion of SS choices in the Willpower task. Improved self-control in the Precommitment task (defined as the difference between the proportion of LL choices in the Precommitment task and the average proportion of LL choices across the other tasks) was positively correlated with impulsivity (Study 1: r = 0.62, p < 0.001; Study 2: r = 0.50, p = 0.020). To identify brain regions involved in the effortful inhibition of impulses, we examined neural activity during the delay period.

To independently test a role for glutamate in the generation of a

To independently test a role for glutamate in the generation of adult rhythms in DD, we misexpressed Gad1, as in larvae ( Figure 5), to reduce presynaptic glutamate. This specifically affects glutamate levels because no adult clock neurons are GABAergic ( Dahdal et al., 2010 and Hamasaka et al., 2005). tim-Gal4; Pdf-Gal80 > Gad1 flies had lower power rhythms than control flies, whereas tim-Gal4; cry-Gal80 > Gad1 flies had robust DD rhythms ( Figures 7D–7F and Table 1). Thus, two independent manipulations of glutamate signaling indicate that glutamate released from CRY+ non-LNv clock neurons is required for

robust locomotor activity rhythms. However, the rhythms of tim > + VGlutRNAi and tim-Gal4; Pdf-Gal80 > Gad1 flies are both stronger AZD2281 cell line than tim-Gal4; Pdf-Gal80

> dORKΔC flies, suggesting that additional signals from non-LNvs contribute to rhythmic behavior. This interpretation makes sense given the diversity of Drosophila adult clock neurons and the incomplete arrhythmicity of even mutants in Pdf, the major circadian neuropeptide ( Renn this website et al., 1999). Taking all of the adult data together, we find evidence that the principles we identified in the larval circadian network may also operate in adult flies. Specifically, our broad manipulations to adult non-LNv clock neurons indicate that non-LNv signals (1) are important for strong adult rhythms, (2) may gate LNv outputs to shape activity at dawn, and (3) include glutamate. We identified some of the network logic Suplatast tosilate that helps generate a simple rhythmic behavior through precise genetic manipulations

of the larval circadian circuit and extended these findings to the more complex adult circadian network. Previous studies have shown that intercellular signaling in clock neuron networks promotes molecular clock synchrony (Lin et al., 2004, Maywood et al., 2006 and Stoleru et al., 2005) and can strengthen genetically weak molecular clocks (Liu et al., 2007). Our study increases the importance of circadian neural networks by finding that non-LNv clock neurons are as important as the “master” pacemaker LNv clock neurons for rhythmic behavior both in larvae and adult flies. However, LNvs can still be considered pacemakers in DD because most manipulations to non-LNv clock neurons do not affect period length. Non-LNv signals appear to gate pacemaker neuron activity. Why is this necessary when LNvs have their own intrinsic excitability rhythms? We propose that the interaction of two oscillators with opposite signs helps reduce the time when LNvs signal. Without signaling from non-LNvs, adult locomotor activity rhythms are weak and activity is distributed throughout the day and night as in tim-Gal4; Pdf-Gal80 > dORKΔC flies. In contrast, in tim-Gal4; Pdf-Gal80 > NaChBac flies, the timing of locomotor activity is narrowed.

, energetic collapse due to total ATP depletion and inability to

, energetic collapse due to total ATP depletion and inability to maintain Ca2+ homeostasis) ( Yao et al., 2011) ( Figure 4D). The ATP consumption rates were estimated by measurement of the energy capacity after inhibition of glycolysis and F1F0-ATP synthase with IAA (100 μM) and oligomycin (0.2 μg/ml), respectively, showing no differences between patient and control fibroblasts

( Figure 4E). However, the ATP production rates in patient fibroblasts monitored by inhibition of glycolysis (IAA, 100 μM) and respiration (NaCN, 1 mM) were found to be significantly decreased compared to controls ( Figure 4F) (energy capacity: patient 1 = 41% ± 6%; patient 2 = 55% ± 8%; patient 3 = 60% ± 6%; control 1 = 100% ± 0%; control 2 = 88% ± 12%; control 3 = 86% ± 8%; n = 3). These results show that VCP-deficient cells Talazoparib manufacturer generate less ATP than control cells but also demonstrate

the vulnerability of these cells selleck products to chemical ischemia ( Figure 4F). As the energy factories of the cells, mitochondria play a vital role in neurons, in which oxidative phosphorylation is the main source of ATP. Previous studies have shown that pathogenic VCP mutations modulate VCP ATPase activity in vitro ( Halawani et al., 2009) and that they are associated with altered cellular ATP levels in Drosophila ( Chan et al., 2012; Chang et al., 2011; Manno et al., 2010). In this study, we investigated the mitochondrial bioenergetics in VCP-deficient cells and in fibroblasts with VCP mutations from IBMPFD patients. We show that loss of VCP function is associated with decreased ΔΨm in the above cell models and in mouse cortical primary neurons and astrocytes. VCP deficiency further results in increased mitochondrial respiration and uncoupling. These observations are accompanied by decreased ATP levels due to lower ATP production. A number of prior studies have observed altered mitochondrial respiratory complex function in ALS disease models including postmortem brain and spinal cord tissue (Bowling et al., 1993; Wiedemann et al., 2002), patient lymphocytes (Ghiasi et al., 2012), and

a transgenic mouse model of ALS (Jung et al., 2002). Despite these findings, there remains some controversy surrounding the dysfunction of mitochondrial respiratory chain complexes in ALS, and we previously found normal activity PIK3C2G in muscle, myoblasts, fibroblasts, and cybrids from patients (Bradley et al., 2009). Accordingly, our results strongly suggest that there is no impairment of mitochondrial respiratory complexes in any of the fibroblasts from the IBMFPD patients carrying the VCP pathogenic mutations. We observed that ΔΨm was decreased in all the VCP-deficient cell models. ΔΨm is a key indicator of mitochondrial viability, as it reflects the pumping of hydrogen ions across the inner membrane during the process of electron transport, the driving force behind ATP production.

, 2010) For immunohistochemical analyses,

mice were deep

, 2010). For immunohistochemical analyses,

mice were deeply anesthetized selleck and transcardially perfused with PBS followed by 4% PFA. Postfixed brains were sectioned coronally at a thickness of 40 μm, followed by permeabilization, blocking, and incubation with primary antibodies overnight. Following incubation with fluorescently tagged secondary antibodies, slices were mounted and imaged. For details, see the Supplemental Experimental Procedures. Transverse hippocampal slices (400 μm) were prepared from 4- to 6-week-old mice of the four genotypes described above. Slice preparation and all LTD experiments were performed as described previously (Sharma et al., 2010). For details see supplementary content. Transverse hippocampal slices (400 μm) were obtained from 4- to 6-week-old mice of the four genotypes. Puromycin labeling of the slices was adapted from procedures described previously (Hoeffer et al., 2011). For details,

see the Supplemental Experimental Procedures. Spine number and morphological analyses were done on rapid Golgi-Cox-stained brain sections using a protocol described previously (Hayashi et al., 2007). For details, see the Supplemental Experimental Procedures. Two independent cohorts of mice of each genotype (a total of 8–12 per genotype) were used for the behavioral tests. Mice were 4–6 months of age, and all mice used were male. The behavioral tests were conducted in increasing order of difficulty and stress ranging from open field analysis, Venetoclax mw marble-burying, rotarod, social interaction, novel object recognition, and Y-maze choice arm reversal. All tests were performed in conditions and in a manner as described

previously (Hoeffer et al., 2008). For details, see the Supplemental Experimental Procedures. This work was supported by NIH grants NS034007 and NS047384 (E.K.), FRAXA Research Foundation (E.K.), and a FRAXA Postdoctoral Fellowship (A.B). J.P.M. was supported by a summer training grant, NSF REU Site Grant in Neural Science DBI 1004172. “
“Robo receptors are important regulators of axon guidance and cell migration in vertebrates and invertebrates (Brose et al., 1999; Dickson and Gilestro, 2006; Legg et al., to 2008). In response to Slit proteins, Robo signaling influences the cytoskeleton to promote repulsion, attraction, or branching, depending on the cellular context (Kidd et al., 1998; Kramer et al., 2001; Long et al., 2004; Wang et al., 1999; Whitford et al., 2002), which allows for a great diversity of biological functions. Using similar mechanisms, Slit/Robo signaling also regulates a large variety of morphogenetic processes outside the central nervous system (CNS), from leukocyte chemotaxis and angiogenesis to kidney and cardiac development (Fish et al., 2011; Grieshammer et al., 2004; Kramer et al., 2001; Legg et al., 2008; London and Li, 2011; Ypsilanti et al., 2010).