Calcrete profile development in quaternary alluvial sequences, southeast Spain: Implications for using calcretes as a basis for landform chronologies

A detailed study of the morphology and micro-morphology of Quaternary alluvial calcrete profiles from the Sorbas Basin shows that calcretes may be morphologically simple or complex. The 'simple' profiles reflect pedogenesis occurring after alluvial terrace formation and consist of a single pedogenic horizon near the land surface. The 'complex' profiles reflect the occurrence of multiple calcrete events during terrace sediment aggradation and further periods of pedogenesis after terrace formation. These 'complex' calcrete profiles are consequently described as composite profiles. The exact morphology of the composite profiles depends upon: (1) the number of calcrete-forming events occurring during terrace sediment aggradation; (2) the amount of sediment accretion that occurs between each period of calcrete formation; and (3) the degree of pedogenesis after terrace formation. Simple calcrete profiles are most useful in establishing landform chronologies because they represent a single phase of pedogenesis after terrace formation. Composite profiles are more problematic. Pedogenic calcretes that form within them may inherit carbonate from calcrete horizons occurring lower down in the terrace sediments. In addition erosion may lead to the exhumation of older calcretes within the terrace sediment. Calcrete 'inheritance' may make pedogenic horizons appear more mature than they actually are and produce horizons containing carbonate embracing a range of ages. Calcrete exhumation exposes calcrete horizons whose morphology and radiometric ages are wholly unrelated to terrace surface age. Composite profiles are, therefore, only suitable for chronological studies if the pedogenic horizon capping the terrace sequence can be clearly distinguished from earlier calcrete-forming events. Thus, a detailed morphological/micro-morphological study is required before any chronological study is undertaken. This is the only way to establish whether particular calcrete profiles are suitable for dating purposes. Copyright (C) 2003 John Wiley Sons, Ltd.

The time-course of morphological constraints: a study of plurals inside derived words

The avoidance of regular but not irregular plurals inside compounds (e.g. *rats eater vs. mice eater) has been one of the most widely studied morphological phenomena in the psycholinguistics literature. To examine whether the constraints that are responsible for this contrast have any general significance beyond compounding, we investigated derived word forms containing regular and irregular plurals in two experiments. Experiment 1 was an offline acceptability judgment task, and experiment 2 measured eye movements during reading derived words containing regular and irregular plurals and uninflected base nouns. The results from both experiments show that the constraint against regular plurals inside compounds generalizes to derived words. We argue that this constraint cannot be reduced to phonological properties, but is instead morphological in nature. The eye-movement data provide detailed information on the time-course of processing derived word forms indicating that early stages of processing are affected by a general constraint that disallows inflected words from feeding derivational processes, and that the more specific constraint against regular plurals comes in at a subsequent later stage of processing. We argue that these results are consistent with stage-based models of language processing.

A phyloclimatic study of Cyclamen

Background: The impact of global climate change on plant distribution, speciation and extinction is of current concern. Examining species climatic preferences via bioclimatic niche modelling is a key tool to study this impact. There is an established link between bioclimatic niche models and phylogenetic diversification. A next step is to examine future distribution predictions from a phylogenetic perspective. We present such a study using Cyclamen (Myrsinaceae), a group which demonstrates morphological and phenological adaptations to its seasonal Mediterranean-type climate. How will the predicted climate change affect future distribution of this popular genus of garden plants? Results: We demonstrate phylogenetic structure for some climatic characteristics, and show that most Cyclamen have distinct climatic niches, with the exception of several wide-ranging, geographically expansive, species. We reconstruct climate preferences for hypothetical ancestral Cyclamen. The ancestral Cyclamen lineage has a preference for the seasonal Mediterranean climate characteristic of dry summers and wet winters. Future bioclimatic niches, based on BIOCLIM and Maxent models, are examined with reference to a future climate scenario for the 2050s. Over the next 50 years we predict a northward shift in the area of climatic suitability, with many areas of current distribution becoming climatically unsuitable. The area of climatic suitability for every Cyclamen species is predicted to decrease. For many species, there may be no areas with a suitable climate regardless of dispersal ability, these species are considered to be at high risk of extinction. This risk is examined from a phylogenetic perspective. Conclusion: Examining bioclimatic niches from a phylogenetic perspective permits novel interpretations of these models. In particular, reconstruction of ancestral niches can provide testable hypothesis about the historical development of lineages. In the future we can expect a northwards shift in climatic suitability for the genus Cyclamen. If this proves to be the case then dispersal is the best chance of survival, which seems highly unlikely for ant-dispersed Cyclamen. Human-assisted establishment of Cyclamen species well outside their native ranges offers hope and could provide the only means of dispersal to potentially suitable future environments. Even without human intervention the phylogenetic perspective demonstrates that major lineages could survive climate change even if many species are lost.

Nomenclature in the dock. Overclassification leads to instability: a case study in the horticulturally important genus Cyclamen (Myrsinaceae)

Nine different classifications have been produced in the last 70 years for the horticulturally valuable genus Cyclamen, a small genus with fewer than 30 species. These classifications, generated by intuitive methods and cladistic analyses, incorporated a total of four infrageneric ranks above that of species and were based on data from morphology, cytology and DNA sequencing. Our results, based on cladistic analyses of three independent data sources − nrDNA ITS, cpDNA trnL intron and morphological data − reveal good resolution only in nrDNA sequence data. However, when these three data sources are combined they provide stronger resolution and support for three major clades, only one of which, subgenus Psilanthum, has been consistently supported in previous classifications. The differing infrageneric classifications produced in Cyclamen result from varying taxon sampling, differing interpretation of morphological data, changes in the sources and analysis of data, and inconsistent application of names. Extensive subdivision of small genera in the absence of adequate data that could provide evidence for consistent patterns of relationship is premature and leads to a proliferation of names.© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146, 339-349.

Quantitative and qualitative differences in morphological traits revealed between diploid Fragaria species

Background and Aims: The aims of this investigation were to highlight the qualitative and quantitative diversity apparent between nine diploid Fragaria species and produce interspecific populations segregating for a large number of morphological characters suitable for quantitative trait loci analysis. Methods: A qualitative comparison of eight described diploid Fragaria species was performed and measurements were taken of 23 morphological traits from 19 accessions including eight described species and one previously undescribed species. A principal components analysis was performed on 14 mathematically unrelated traits from these accessions, which partitioned the species accessions into distinct morphological groups. Interspecific crosses were performed with accessions of species that displayed significant quantitative divergence and, from these, populations that should segregate for a range of quantitative traits were raised. Key Results: Significant differences between species were observed for all 23 morphological traits quantified and three distinct groups of species accessions were observed after the principal components analysis. Interspecific crosses were performed between these groups, and F2 and backcross populations were raised that should segregate for a range of morphological characters. In addition, the study highlighted a number of distinctive morphological characters in many of the species studied. Conclusions: Diploid Fragaria species are morphologically diverse, yet remain highly interfertile, making the group an ideal model for the study of the genetic basis of phenotypic differences between species through map-based investigation using quantitative trait loci. The segregating interspecific populations raised will be ideal for such investigations and could also provide insights into the nature and extent of genome evolution within this group.

A genetic linkage map of microsatellite, gene-specific and morphological markers in diploid Fragaria

Diploid Fragaria provide a potential model for genomic studies in the Rosaceae. To develop a genetic linkage map of diploid Fragaria, we scored 78 markers (68 microsatellites, one sequence-characterised amplified region, six gene-specific markers and three morphological traits) in an interspecific F2 population of 94 plants generated from a cross of F.vesca f. semperflorens × F. nubicola. Co-segregation analysis arranged 76 markers into seven discrete linkage groups covering 448 cM, with linkage group sizes ranging from 100.3 cM to 22.9 cM. Marker coverage was generally good; however some clustering of markers was observed on six of the seven linkage groups. Segregation distortion was observed at a high proportion of loci (54%), which could reflect the interspecific nature of the progeny and, in some cases, the self-incompatibility of F. nubicola. Such distortion may also account for some of the marker clustering observed in the map. One of the morphological markers, pale-green leaf (pg) has not previously been mapped in Fragaria and was located to the mid-point of linkage group VI. The transferable nature of the markers used in this study means that the map will be ideal for use as a framework for additional marker incorporation aimed at enhancing and resolving map coverage of the diploid Fragaria genome. The map also provides a sound basis for linkage map transfer to the cultivated octoploid strawberry.

Effects of dendritic morphology on CA3 pyramidal cell electrophysiology: a simulation study

We investigated the effect of morphological differences on neuronal firing behavior within the hippocampal CA3 pyramidal cell family by using three-dimensional reconstructions of dendritic morphology in computational simulations of electrophysiology. In this paper, we report for the first time that differences in dendritic structure within the same morphological class can have a dramatic influence on the firing rate and firing mode (spiking versus bursting and type of bursting). Our method consisted of converting morphological measurements from three-dimensional neuroanatomical data of CA3 pyramidal cells into a computational simulator format. In the simulation, active channels were distributed evenly across the cells so that the electrophysiological differences observed in the neurons would only be due to morphological differences. We found that differences in the size of the dendritic tree of CA3 pyramidal cells had a significant qualitative and quantitative effect on the electrophysiological response. Cells with larger dendritic trees: (1) had a lower burst rate, but a higher spike rate within a burst, (2) had higher thresholds for transitions from quiescent to bursting and from bursting to regular spiking and (3) tended to burst with a plateau. Dendritic tree size alone did not account for all the differences in electrophysiological responses. Differences in apical branching, such as the distribution of branch points and terminations per branch order, appear to effect the duration of a burst. These results highlight the importance of considering the contribution of morphology in electrophysiological and simulation studies.

Faunal evidence of a Holocene pluvial phase in Southern Arabia with remarks on the morphological variability of Helenina anderseni

Although foraminifera have been found living in inland saline lakes isolated from the sea, this phenomenon has rarely been recognized in the fossil record. This study documents the occurrence of benthic foraminifera in Holocene lake sediments located nearly 500 km inland from the Red Sea, in the Al-Mundafan region of southern Saudi Arabia. The lake formed during a regional pluvial period, 10,500–6000 yr BP. The presence of foraminifera and brackish charophytes in the studied section represent an interval when the lake was slightly brackish due to high evaporation. The studied sediments yielded a bispecific benthic foraminiferal fauna comprised of Helenina anderseni and Trichohyalus aguayoi, as well as the brackish charophyte genus Lamprothamnium. The benthic foraminifera are species characteristic of mangrove swamps, salt marshes, and lagoons, which are environments currently widespread along the Red Sea coasts. Because the Al Mundafan area was never connected to the sea during the Quaternary, wading birds must have been the vector that transported the foraminifera to the paleolake.

A study of the microstructure and condition of thin sheets of painting support ivory using SEM and ESEM

This paper outlines a study of the microstructure of thin sheets of ivory used as a painting support for portrait miniatures. Warping of the ivory support is one of the main problems commonly found in portrait miniatures from the late eighteenth century and early nineteenth century. Portrait miniatures from this period are painted on very thin sheets of ivory that are often only 0.2 mm in thickness. Warping can lead to cracking of the ivory and can also accentuate flaking of the paint layer. The problem of warping in ivory has thus been of long-term interest to conservators who deal with portrait miniatures, including those at the Victoria and Albert (V&A) Museum. The causes of warping are complex. However, it should be noted that artists normally stuck the thin ivory sheets onto paper or card before commencing the painting. The possible causes of warping therefore are thought to relate to the differential reactions of the ivory/adhesive/paper or card layers to changes in relative humidity (RH). It is well known that ivory is hygroscopic and anisotropic.1 However, only a few scientific studies have been carried out related to this subject and systematic analyses of the morphological and microstructural changes due to changes in RH or moisture in such thin sheets of ivory have yet to be investigated.

A molecular, morphological and cytological investigation of the identity of non-native Ludwigia (Onagraceae) populations in Britain

A combined molecular, morphological and cytological analysis was used to study the identity and number of species of Ludwigia section Oligospermum occurring in British waterways. Only one taxon was identified for which the name L. grandiflora subsp. hexapetala (Hook. & Arn.) G.L. Nesom & Kartesz is preferred. A chromosome count of 2n = 80 was made for all plants tested and DNA evidence demonstrates that at least two clones are present in Britain. Morphological characters to differentiate L.grandiflora subsp. hexapetala and L. peploides subsp. montevidensis (Spreng.) P.H. Raven are provided. Though the production of fruit in Britain by apparently isolated populations is noted, repeated introduction into the wild from gardens is judged to be primarily responsible for the British distribution of the taxon. Legislative implications are considered and an amendment to Schedule 9 of the UK Wildlife and Countryside Act (1981) is strongly advocated.

The time-course of morphological constraints: Evidence from eye-movements during reading

Lexical compounds in English are constrained in that the non-head noun can be an irregular but not a regular plural (e.g. mice eater vs. *rats eater), a contrast that has been argued to derive from a morphological constraint on modifiers inside compounds. In addition, bare nouns are preferred over plural forms inside compounds (e.g. mouse eater vs. mice eater), a contrast that has been ascribed to the semantics of compounds. Measuring eyemovements during reading, this study examined how morphological and semantic information become available over time during the processing of a compound. We found that the morphological constraint affected both early and late eye-movement measures, whereas the semantic constraint for singular non-heads only affected late measures of processing. These results indicate that morphological information becomes available earlier than semantic information during the processing of compounds.

The time-course of morphological processing in a second language

We report findings from psycholinguistic experiments investigating the detailed timing of processing morphologically complex words by proficient adult second (L2) language learners of English in comparison to adult native (L1) speakers of English. The first study employed the masked priming technique to investigate -ed forms with a group of advanced Arabic-speaking learners of English. The results replicate previously found L1/L2 differences in morphological priming, even though in the present experiment an extra temporal delay was offered after the presentation of the prime words. The second study examined the timing of constraints against inflected forms inside derived words in English using the eye-movement monitoring technique and an additional acceptability judgment task with highly advanced Dutch L2 learners of English in comparison to adult L1 English controls. Whilst offline the L2 learners performed native-like, the eye-movement data showed that their online processing was not affected by the morphological constraint against regular plurals inside derived words in the same way as in native speakers. Taken together, these findings indicate that L2 learners are not just slower than native speakers in processing morphologically complex words, but that the L2 comprehension system employs real-time grammatical analysis (in this case, morphological information) less than the L1 system.

Morphological decomposition in Broca's aphasia

Background: Few studies have investigated how individuals diagnosed with post-stroke Broca’s aphasia decompose words into their constituent morphemes in real-time processing. Previous research has focused on morphologically complex words in non-time-constrained settings or in syntactic frames, but not in the lexicon. Aims: We examined real-time processing of morphologically complex words in a group of five Greek-speaking individuals with Broca’s aphasia to determine: (1) whether their morphological decomposition mechanisms are sensitive to lexical (orthography and frequency) vs. morphological (stem-suffix combinatory features) factors during visual word recognition, (2) whether these mechanisms are different in inflected vs. derived forms during lexical access, and (3) whether there is a preferred unit of lexical access (syllables vs. morphemes) for inflected vs. derived forms. Methods & Procedures: The study included two real-time experiments. The first was a semantic judgment task necessitating participants’ categorical judgments for high- and low-frequency inflected real words and pseudohomophones of the real words created by either an orthographic error at the stem or a homophonous (but incorrect) inflectional suffix. The second experiment was a letter-priming task at the syllabic or morphemic boundary of morphologically transparent inflected and derived words whose stems and suffixes were matched for length, lemma and surface frequency. Outcomes & Results: The majority of the individuals with Broca’s aphasia were sensitive to lexical frequency and stem orthography, while ignoring the morphological combinatory information encoded in the inflectional suffix that control participants were sensitive to. The letter-priming task, on the other hand, showed that individuals with aphasia—in contrast to controls—showed preferences with regard to the unit of lexical access, i.e., they were overall faster on syllabically than morphemically parsed words and their morphological decomposition mechanisms for inflected and derived forms were modulated by the unit of lexical access. Conclusions: Our results show that in morphological processing, Greek-speaking persons with aphasia rely mainly on stem access and thus are only sensitive to orthographic violations of the stem morphemes, but not to illegal morphological combinations of stems and suffixes. This possibly indicates an intact orthographic lexicon but deficient morphological decomposition mechanisms, possibly stemming from an underspecification of inflectional suffixes in the participants’ grammar. Syllabic information, however, appears to facilitate lexical access and elicits repair mechanisms that compensate for deviant morphological parsing procedures.