The neural development and organization of letter recognition: Evidence from functional neuroimaging, computational modeling, and behavioral studies

Although much of the brain’s functional organization is genetically predetermined, it appears that some noninnate functions can come to depend on dedicated and segregated neural tissue. In this paper, we describe a series of experiments that have investigated the neural development and organization of one such noninnate function: letter recognition. Functional neuroimaging demonstrates that letter and digit recognition depend on different neural substrates in some literate adults. How could the processing of two stimulus categories that are distinguished solely by cultural conventions become segregated in the brain? One possibility is that correlation-based learning in the brain leads to a spatial organization in cortex that reflects the temporal and spatial clustering of letters with letters in the environment. Simulations confirm that environmental co-occurrence does indeed lead to spatial localization in a neural network that uses correlation-based learning. Furthermore, behavioral studies confirm one critical prediction of this co-occurrence hypothesis, namely, that subjects exposed to a visual environment in which letters and digits occur together rather than separately (postal workers who process letters and digits together in Canadian postal codes) do indeed show less behavioral evidence for segregated letter and digit processing.

Tomato Fructokinases Exhibit Differential Expression and Substrate Regulation1

Two divergent genes encoding fructokinase, Frk1 and Frk2, have been previously shown to be expressed in tomato (Lycopersicon esculentum L.) and have now been further characterized with regard to their spatial expression and the enzymic properties of the encoded proteins. Frk1 and Frk2 mRNA levels were coordinately induced by exogenous sugar, indicating that both belong to the growing class of sugar-regulated genes. However, in situ hybridization indicated that Frk1 and Frk2 were expressed in a spatially distinct manner, with Frk2 mRNA primarily localized in cells of the fruit pericarp, which store starch, and Frk1 mRNA distributed ubiquitously in pericarp tissue. To evaluate the biochemical characteristics of the products of the Frk1 and Frk2 genes, each cDNA was expressed in a mutant yeast (Saccharomyces cerevisiae) line defective in hexose phosphorylation and unable to grow on glucose or fructose (Fru). Both Frk1 and Frk2 proteins expressed in yeast conferred the ability to grow on Fru and exhibited fructokinase activity in vitro. Although both Frk1 and Frk2 both utilized Fru as a substrate, only Frk2 activity was inhibited at high Fru concentrations. These results indicate that Frk2 can be distinguished from Frk1 by its sensitivity to substrate inhibition and by its temporal and spatial pattern of expression, which suggests that it plays a primary role in plant cells specialized for starch storage.

Three 1-Aminocyclopropane-1-Carboxylate Synthase Genes Regulated by Primary and Secondary Pollination Signals in Orchid Flowers1

The temporal and spatial expression patterns of three 1-aminocyclopropane-1-carboxylate (ACC) synthase genes were investigated in pollinated orchid (Phalaenopsis spp.) flowers. Pollination signals initiate a cascade of development events in multiple floral organs, including the induction of ethylene biosynthesis, which coordinates several postpollination developmental responses. The initiation and propagation of ethylene biosynthesis is regulated by the coordinated expression of three distinct ACC synthase genes in orchid flowers. One ACC synthase gene (Phal-ACS1) is regulated by ethylene and participates in amplification and interorgan transmission of the pollination signal, as we have previously described in a related orchid genus. Two additional ACC synthase genes (Phal-ACS2 and Phal-ACS3) are expressed primarily in the stigma and ovary of pollinated orchid flowers. Phal-ACS2 mRNA accumulated in the stigma within 1 h after pollination, whereas Phal-ACS1 mRNA was not detected until 6 h after pollination. Similar to the expression of Phal-ACS2, the Phal-ACS3 gene was expressed within 2 h after pollination in the ovary. Exogenous application of auxin, but not ACC, mimicked pollination by stimulating a rapid increase in ACC synthase activity in the stigma and ovary and inducing Phal-ACS2 and Phal-ACS3 mRNA accumulation in the stigma and ovary, respectively. These results provide the basis for an expanded model of interorgan regulation of three ACC synthase genes that respond to both primary (Phal-ACS2 and Phal-ACS3) and secondary (Phal-ACS1) pollination signals.

Cell cycling and patterned cell proliferation in the Drosophila wing during metamorphosis.

In metamorphosing wing discs, progression through the cell cycle takes place, as in larval discs, in nonclonally derived clusters of cells synchronized in the same cell cycle stage. Contrary to early discs, there are temporal and spatial heterogeneities in cell proliferation associated with wing margin, vein, intervein, and middle intervein territories. Within these territories, there are no indications of a wave progression of the cell cycle. Mitotic orientations are, as in early discs, at random but there is a preferential allocation of postmitotic cells along the proximodistal axis, thus explaining the elongated shape of the resulting clones along this axis. Shapes of clones in mature discs and in evaginated wings are similar, thus excluding major morphogenetic movements during evagination. After the proliferative period, all the cells are arrested in G1 phase. The final number of cells of the wing is fixed independently of experimental perturbations that alter the cell division schedule. These results are discussed in the context of a model of wing morphogenesis.

Modular cis-regulatory organization of developmentally expressed genes: two genes transcribed territorially in the sea urchin embryo, and additional examples.

The cis-regulatory systems that control developmental expression of two sea urchin genes have been subjected to detailed functional analysis. Both systems are modular in organization: specific, separable fragments of the cis-regulatory DNA each containing multiple transcription factor target sites execute particular regulatory subfunctions when associated with reporter genes and introduced into the embryo. The studies summarized here were carried out on the CyIIIa gene, expressed in the embryonic aboral ectoderm and on the Endo16 gene, expressed in the embryonic vegetal plate, archenteron, and then midgut. The regulatory systems of both genes include modules that control particular aspects of temporal and spatial expression, and in both the territorial boundaries of expression depend on a combination of negative and positive functions. In both genes different regulatory modules control early and late embryonic expression. Modular cis-regulatory organization is widespread in developmentally regulated genes, and we present a tabular summary that includes many examples from mouse and Drosophila. We regard cis-regulatory modules as units of developmental transcription control, and also of evolution, in the assembly of transcription control systems.

Origin and evolution of circular waves and spirals in Dictyostelium discoideum territories.

Randomly distributed Dictyostelium discoideum cells form cooperative territories by signaling to each other with cAMP. Cells initiate the process by sending out pulsatile signals, which propagate as waves. With time, circular and spiral patterns form. We show that by adding spatial and temporal noise to the levels of an important regulator of external cAMP levels, the cAMP phosphodiesterase inhibitor, we can explain the natural progression of the system from randomly firing cells to circular waves whose symmetries break to form double- and single- or multi-armed spirals. When phosphodiesterase inhibitor is increased with time, mimicking experimental data, the wavelength of the spirals shortens, and a proportion of them evolve into pairs of connected spirals. We compare these results to recent experiments, finding that the temporal and spatial correspondence between experiment and model is very close.

Temporal and molecular characteristics of mutations induced by ethylnitrosourea in germ cells isolated from seminiferous tubules and in spermatozoa of lacZ transgenic mice.

The lacZ transgenic mouse (Muta mouse) model was used to examine the timing of ethylnitrosourea (ENU)-induced mutations in germ cells. The spectrum of mutations was also determined. Animals received five daily treatments with ENU at 50 mg/kg and were sampled at times up to 55 days after treatment. In mixed germ-cell populations isolated from seminiferous tubules, there was little increase in the mutant frequency 5 days after treatment; subsequently, there was a continuous increase until the maximum (17.5-fold above background) was reached by approximately 35 days. In the spermatozoa, an increase in mutant frequency was not seen until 20 days after treatment, with the maximum (4.3-fold above background) being achieved no sooner than approximately 35 days. Based on the timing of sampling, these data demonstrate the detection of both spermatogonial and postspermatogonial, mutations. The most prominent feature of the ENU-induced base-pair mutations in testicular germ cells sampled 55 days after treatment is that 70% are induced in A.T base pairs, compared to only 16% in spontaneous mutations. These findings are consistent with comparable data from ENU studies using assays for inherited germ-cell mutations in mice. This study has demonstrated the utility and potential of the transgenic mouse lacZ model (Muta mouse) for the detection and study of germ-cell mutations and provides guidance in the selection of simplified treatment and sampling protocols.

Primary structure of hepatocyte nuclear factor/forkhead homologue 4 and characterization of gene expression in the developing respiratory and reproductive epithelium.

Members of the winged helix/forkhead family of transcription factors are believed to play a role in cell-specific gene expression. A cDNA encoding a member of this family of proteins, termed hepatocyte nuclear factor/forkhead homologue 4 (HFH-4), has been isolated from rat lung and rat testis cDNA libraries. This cDNA contains an open reading frame of 421 amino acids with a conserved DNA binding domain and several potential transactivating regions. During murine lung development, a single species of HFH-4-specific transcript (2.4 kb long) is first detected precisely at the start of the late pseudoglandular stage (embryonic day 14.5) and, by in situ hybridization, is specifically localized to the proximal pulmonary epithelium. The unique temporal and spatial pattern of HFH-4 gene expression in the developing lung defines this protein as a marker for the initiation of bronchial epithelial cell differentiation and suggests that it may play an important role in cell fate determination during lung development. In addition to expression in the pulmonary epithelium, RNA blot analysis reveals 2.4-kb HFH-4 transcripts in the testis and oviduct. By using mice with genetic defects in spermatogenesis, HFH-4 expression in the testis is found to be associated with the appearance of haploid germ cells and in situ hybridization studies indicate that HFH-4 expression is confined to stages I-VII of spermatogenesis. This pattern of HFH-4 gene expression during the early stages of differentiation of haploid germ cells suggests that HFH-4 may play a role in regulating stage-specific gene expression and cell-fate determination during lung development and in spermatogenesis.

The episcopal complex of Eio-El Tolmo de Minateda (Hellín, Albacete, Spain). Architecture and spatial organization. 7th to 8th centuries AD

The episcopal complex of Eio, located in El Tolmo de Minateda, was built between the end of the 6th century and the beginning of the 7th century, possibly as a political decision taken by the ecclesiastical authority in the capital of the Visigothic kingdom (Toletum). With the comprehensive study of the whole complex presented below (construction cycles, furniture, decoration and location of spaces), we can interpret the function of each space in the basilica and the domus episcopi, the liturgical and general movement routes, the existence of some hierarchical environments, and specify the chronological development of the buildings. After the Arab-Berber conquest of Hispania in the early 8th century, the whole complex will experience a series of transformations that will convert the religious and monumental public area into a private, residential and industrial Islamic quarter.

Evaluating Taphonomic Bias in a Storm-Disturbed Carbonate Platform: Effects of Compositional and Environmental Factors in Lower Jurassic Brachiopod Accumulations (Eastern Subbetic Basin, Spain)

In order to evaluate taxonomic and environmental control on the preservation pattern of brachiopod accumulations, sedimentologic and taphonomic data have been integrated with those inferred from the structure of brachiopod accumulations from the easternmost Lower Jurassic Subbetic deposits in Spain. Two brachiopod communities (Praesphaeroidothyris and Securina communities) were distinguished showing a mainly free-lying way of life in soft-bottom habitats. Three taphofacies are discriminated based on proportion of disarticulation, fragmentation, packing, and shell filling. Taphofacies 1 is represented by thinly fragmented, dispersed brachiopod shells in wackestone beds. Taphofacies 2 is spatially restricted to small lenses where shells are poorly fragmented, rarely disarticulated, usually void filled, and highly packed. Taphofacies 3 is represented by mud or cement filled, loosely packed, articulated brachiopods forming large pocket-like structures. Temporal and spatial averaging were minimally involved in taphofacies 2 and 3. It is interpreted that patchy preservation implies preservation of primary original patchiness of brachiopod communities on the seafloor. The origin of shell-rich taphofacies (2 and 3) is related to rapid burial due to episodic storm activity, while shell-poor taphofacies 1 records background conditions. The nature and comparative diversity of these taphofacies underscores the importance of rapid burial for shell beds preservation. Differences in preservation between taphofacies 2 and 3 are mainly related to environmental criteria, most importantly storm energy and water depth. In contrast, the taxonomic-specific pattern of the communities is a subordinate element of control, controlling only minor within-taphofacies differences in preservation.

(Table 2) Radiocarbon ages on Arctic box and multicores

Paleo-sea-ice history in the Arctic Ocean was reconstructed using the sea-ice dwelling ostracode Acetabulastoma arcticum from late Quaternary sediments from the Mendeleyev, Lomonosov, and Gakkel Ridges, the Morris Jesup Rise and the Yermak Plateau. Results suggest intermittently high levels of perennial sea ice in the central Arctic Ocean during Marine Isotope Stage (MIS) 3 (25-45 ka), minimal sea ice during the last deglacial (16-11 ka) and early Holocene thermal maximum (11-5 ka) and increasing sea ice during the mid-to-late Holocene (5-0 ka). Sediment core records from the Iceland and Rockall Plateaus show that perennial sea ice existed in these regions only during glacial intervals MIS 2, 4, and 6. These results show that sea ice exhibits complex temporal and spatial variability during different climatic regimes and that the development of modern perennial sea ice may be a relatively recent phenomenon.

Sr and Nd composition, stable oxygen isotope record and lithogenic flux in eastern Mediterranean sediments

Isotopic ratios of Sr and Nd from lithogenic components of three isochronous core sections recovered from an east-west transect in the Eastern Mediterranean Sea (EMS) have been analyzed. The data are used for a quantitative estimate of the temporal and spatial variation of detrital flux to the EMS, assuming Saharan dust and Aegean/Nile particulate matter as dominant end members. It was established that the carbonate-free Saharan dust flux during deposition of the nonsapropel layers of marine oxygen isotope stage 5.4 (MIS 5.4) was similar to the present flux. During the deposition of sapropels S5 and S6, however, the Saharan dust input was drastically reduced and was not balanced by a change in the riverine influx at this time. Denser vegetation cover during more humid conditions may have reduced physical erosion and sediment removal in the source area. During marine oxygen isotope stage 6.2 (MIS 6.2) a pronounced increase of Saharan dust and detrital influx from the Aegean region is evident and implies more arid conditions in the southern and northern catchment areas. During this period, intersite variations are interpreted in terms of their geographic location relative to the seaways connecting the Aegean Sea and EMS. The width of the straits and hence the amount of sediment entering the eastern basins may have been affected by a low sea level that impeded interbasin sediment dispersal.

Tree line shifts and disturbance events for sites studied in the Torneträsk area of sub-Arctic Sweden for the period 1800-2009

Aim: Models project that climate warming will cause the tree line to move to higher elevations in alpine areas and more northerly latitudes in Arctic environments. We aimed to document changes or stability of the tree line in a sub-Arctic model area at different temporal and spatial scales, and particularly to clarify the ambiguity that currently exists about tree line dynamics and their causes. Location: The study was conducted in the Tornetrask area in northern Sweden where climate warmed by 2.5 °C between 1913 and 2006. Mountain birch (Betula pubescens ssp. czerepanovii) sets the alpine tree line. Methods: We used repeat photography, dendrochronological analysis, field observations along elevational transects and historical documents to study tree line dynamics. Results: Since 1912, only four out of eight tree line sites had advanced: on average the tree line had shifted 24 m upslope (+0.2 m/year assuming linear shifts). Maximum tree line advance was +145 m (+1.5 m/year in elevation and +2.7 m/year in actual distance), whereas maximum retreat was 120 m downslope. Counter-intuitively, tree line advance was most pronounced during the cooler late 1960s and 1970s. Tree establishment and tree line advance were significantly correlated with periods of low reindeer (Rangifer tarandus) population numbers. A decreased anthropozoogenic impact since the early 20th century was found to be the main factor shaping the current tree line ecotone and its dynamics. In addition, episodic disturbances by moth outbreaks and geomorphological processes resulted in descent and long-term stability of the tree line position, respectively. Main conclusions: In contrast to what is generally stated in the literature, this study shows that in a period of climate warming, disturbance may not only determine when tree line advance will occur but if tree line advance will occur at all. In the case of non-climatic climax tree lines, such as those in our study area, both climate-driven model projections of future tree line positions and the use of the tree line position for bioclimatic monitoring should be used with caution.

Sea surface temperature and stable isotopes from 9 cores of the Northeastern Atlantic

Temporal and spatial patterns in eastern North Atlantic sea-surface temperatures (SST) were reconstructed for marine isotope stage (MIS) 11c using a submeridional transect of five sediment cores. The SST reconstructions are based on planktic foraminiferal abundances and alkenone indices, and are supported by benthic and planktic stable isotope measurements, as well as by ice-rafted debris content in polar and middle latitudes. Additionally, the larger-scale dynamics of the precipitation regime over northern Africa and the western Mediterranean region was evaluated from iron concentrations in marine sediments off NW Africa and planktic d13C in combination with analysis of planktic foraminiferal abundances down to the species level in the Mediterranean Sea. Compared to the modern situation, it is revealed that during entire MIS 11c sensu stricto (ss), i.e., between 420 and 398 ka according to our age models, a cold SST anomaly in the Nordic seas co-existed with a warm SST anomaly in the middle latitudes and the subtropics, resulting in steeper meridional SST gradients than during the Holocene. Such a SST pattern correlates well with a prevalence of a negative mode of the modern North Atlantic Oscillation. We suggest that our scenario might partly explain the longer duration of wet conditions in the northern Africa during MIS 11c compared to the Holocene.

Traffic and spatial impacts and the classification of small highway-bypassed cities.

Texas Department of Transportation, Austin