Reassessing domestication events in the Near East: Einkorn and Triticum urartu

To reassess domestication events in the Near East, accessions of Triticum urartu from a well-described sampling were combined with a representative sample covering the Karacada? Einkorn wheat domestication. The observed DNA separation between the two wheat species accounts for the main differentiation, but geographic variation within T. urartu is evident and so is the domestication scenario among wild, feral, and domesticated Einkorn. In contrast to the clear DNA differences, it is difficult to separate living T. urartu from wild Einkorn based on morphology. With archaeobotanical material a distinction of carbonized remains of these two wheats is considered to be impossible. We reviewed the differences concerning morphology and maturity and combined these observations with information about archaeological sites in the Near East. In conclusion, the excavation sites in the middle Euphrates may contain T. urartu rather than Einkorn wheat and T. urartu may underlie the reported occurrence of the extinct 2-grained domesticated "Einkorn" wheat. The first Einkorn wheat domestication sensu stricto seems to have happened around the Karacada?, as reported earlier. The human dimension shown by the excavation of G?bekli Tepe can explain why domesticated phenotypes might have spread quickly.     

Times of Plenty, Times of Less: Last-Millennium Societal Disruption in the Pacific Basin

During the last millennium in the Pacific Basin (islands and continental rim) there was a marked contrast between ‘times of plenty’ and ‘times of less’ for its human societies. This contrast is attributable to climate and sea-level variations, notably the Medieval Warm Period (a.d. 700–1250) and the Little Ice Age (a.d. 1350–1800) separated by a time of rapid cooling and sea-level fall called the ‘a.d. 1300 Event.’ Outlines of the times of plenty during the Medieval Warm Period and the times of less during the Little Ice Age are given, supported by a number of examples. These confirm a general picture of societal collapse as a result of the a.d. 1300 Event. Well-dated human responses to the a.d. 1300 Event (establishment of fortified settlements, end of ocean voyaging) allow links to potential nonhuman causes to be strengthened. Although more data referring to both (natural) changes and their human effects are needed, a conclusion involving environmental determinism is inescapable.     

Vegetation responses to rapid climatic changes during the last deglaciation 13,500–8,000?years ago on southwest And?ya, arctic Norway

The late-glacial vegetation development in northern Norway in response to climate changes during the Aller?d, Younger Dryas (YD), and the transition to the Holocene is poorly known. Here we present a high-resolution record of floral and vegetation changes at lake Lusvatnet, south-west And?ya, between 13500 and 8000 cal b.p. Plant macrofossil and pollen analyses were done on the same sediment core and the proxy records follow each other very closely. The core has also been analyzed using an ITRAX XRF scanner in order to check the sediment sequence for disturbances or hiatuses. The core has a good radiocarbon-based chronology. The Saksunarvatn tephra fits very well chronostratigraphically. During both the Aller?d and the Younger Dryas time-periods arctic vegetation prevailed, dominated by Salix polaris associated with many typically arctic herbs such as Saxifraga cespitosa, Saxifraga rivularis and Oxyria digyna. Both periods were cold and dry. Between 12450 and 12250 cal b.p. during the Younger Dryas chronozone, the assemblage changed, particularly in the increased abundance of Papaver sect. Scapiflora and other high-Arctic herbs, suggesting the development of polar desert vegetation mainly as a response to increased aridity. After 11520 cal b.p. a gradually warmer and more oceanic climate initiated a succession to dwarf-shrub vegetation and the establishment of Betula woodland after 1,000 years at c. 10520 cal b.p. The overall late-glacial aridity contrasts with oceanic conditions in southern Norway and is probably related to sea-ice extent.     

Genetic diversity of globe artichoke landraces from Sicilian small-holdings: implications for evolution and domestication of the species

Globe artichoke (Cynara cardunculus var. scolymus) is native to the Mediterranean Basin, where it grows in close proximity with its ancestor wild cardoon (C. cardunculus var. sylvestris); its commercial production is mainly based on vegetatively propagated clones which guarantee high yields of marketable product (i.e. immature inflorescence or capitula). A collection of 24 landraces of globe artichoke was made from small-holdings in Sicily, which is assumed to be one of the possible centres of its domestication. These landraces have been cultivated for centuries by local farmers, mainly due to their culinary uniqueness. The collection was characterised for a combination of morphological traits and AFLP, gSSR and cpSSr markers. Molecular analyses included genotypes of wild cardoon collected from different sites in Sicily as well as accessions of the most widely grown Sicilian varietal types: the spiny ‘Spinoso di Palermo’ and the non-spiny ‘Violetto di Sicilia’. The landraces follow a gradient of ‘ennoblement’ towards either the domesticated spiny or the non-spiny types. ‘Cimiciusa di Mazzarino’ was an outlier, in that it resembled the cultivated forms with respect to its AFLP fingerprint, but was more closely related to the wild cardoon on the basis of SSR profile. This particular landrace presents an example of an intermediary form in the domestication process, although it could also have derived from introgression from sympatric wild cardoon, followed by farmer selection. The abundant genetic variation present demonstrates the key role of farmers’ practice in the maintenance of genetic diversity, which should be preserved because of its potential value for plant breeders.     

Critique of ‘Australopithecus afarensis’ as a single species based on dental metrics and morphology

Leonard andHegmon (1987) compare a series of dental metrics of ‘Australopithecus afarensis Johanson, White, andCoppens, 1978’ with criteria for modern apes, to test the hypothesis that ‘A. afarensis’ represents a single species. They also compare the morphology of the lower third premolar. The dental breadth of ‘A. afarensis’ shows a wide range of variation, particularly in the lower third premolar morphology which displays greater variation than in modern apes—yet the study concludes that the single species hypothesis cannot be rejected. The study is flawed by applying criteria for pongids inappropriate for a hominid. When ‘A. afarensis’ is compared with criteria for hominids, the range of variation in dental size, breadth, and third premolar morphology is greater than that in any hominid species. The single species hypothesis is, therefore, once again rejected. Moreover, the name ‘A. afarensis’ is preoccupied byPraeanthropus africanus (Weinert) and must be dropped.     

The distribution, natural habitats and availability of wild cereals in relation to their domestication in the Near East: multiple events, multiple centres

In this article we examine the natural habitats and distribution of the six wild cereals: Triticum urartu (wild urartu wheat), T. boeoticum aegilopoides (single-grained wild einkorn), T. boeoticum thaoudar (two-grained wild einkorn), T. dicoccoides (wild emmer wheat), Secale spp. (wild ryes) and Hordeum spontaneum (wild barley). A comparison of late Pleistocene/early Holocene archaeobotanical assemblages in the Near East with present-day distributions of wild cereals shows a good correlation. The regional variation in the archaeobotanical cereal assemblages and the ensuing domestication provide evidence that different cereal species were domesticated independently in different areas. Some sites were not situated near wild cereal habitats and a few were located outside the limits of distribution, even accounting for moister climatic conditions. I argue here that current models which try to explain the shift to farming have tended to over-emphasize the effect of the Younger Dryas climatic change. First, it would have had only a minor effect on cereal availability. Secondly, agriculture appears to have been established after the Younger Dryas. Thirdly, there is no evidence for a single centre of origin; agriculture arose in widely separated geographic and climatic regions. And fourthly, agriculture depends on stable climatic conditions which were not established until after the Younger Dryas.     

Application of the equilibrium concept to the development of agaric fruit-bodies, with special reference to their straight downward growth in light from below

Equilibrium, a concept of dynamics, is found to be applicable to the phototropic and gravitropic growth in agaric fruit-bodies. The fruit-bodies exposed to light from below grow straight downward without bending upward, and those exposed to light from obliquely below grow first downward and then upward by negative gravitropism. The fruit-bodies exposed to light from above grow upward. Fruit-bodies growing straight downward or upward do not change the direction of growth; they are in ‘equilibria’. The straight downward growth can be regarded as an ‘unstable equilibrium’ having a higher potential, and the straight upward growth as a ‘stable equilibrium’ having a lower potential. The change in the direction of growth can be explained by the change in the potential; the upward bending in fruit-bodies that have grown obliquely downward can be regarded as a ‘transition’ from the unstable equilibrium to the stable one.     

Vegetation development and human occupation in the Damascus region of southwestern Syria from the Late Pleistocene to Holocene

Rockshelter Baaz in the Damascus region of Syria provided a variety of botanical remains from the Late Pleistocene and Early Holocene period. These remains provide new information about the vegetation evolution in this region. The earliest occupational levels correspond with a moisture peak during the Late Pleistocene, between ca. 34–32 kyr b.p., when pine expanded. The next occupations took place during extreme arid conditions, ca. 23–21 kyr b.p., and probably during the Last Glacial Maximum when a steppe vegetation was established. The occupation level of the Younger Dryas, represented by Natufian remains, suggests that the area had been covered by almond-pistachio steppe, similar to later periods of the Early Holocene, and was probably located just outside the range of dense wild cereal stands. There is no drastic impact of the Younger Dryas visible on the vegetation in the botanical remains. The lack of fruits and seeds at Baaz indicates that the site was more likely to have been a temporary hunting post rather than a plant processing site for much of its history. It is ideally suited to this purpose because of its location over the Jaba′deen Pass and the associated springs. However, archaeological remains from the Natufian period, suggest that the site was more permanently occupied during this time. A. Dodonov deceased.     

A 12 000-year record of environmental change in the Lomond Hills, Fife, Scotland: Vegetational and climatic variability

The Lomond Hills of Fife, an isolated upland area rising to over 500 m, provide an opportunity to investigate the effect of altitude on vegetation and climate in an area otherwise dominated by lower-lying land. The West Lomond site contains sediments of the Devensian Late-glacial period; they reveal a well-defined sequence of Bolling-Older Dryas-Allerod-Younger Dryas events, commencing ca. 12 190 radiocarbon years B.P. and a probable Amphi-Atlantic Oscillation between ca. 11 040 and 10 800 B.P. The Holocene record is constrained by low sediment input but does reveal a woodland presence at this altitude, dominated byBetula andCorylus. Size statistics forBetula pollen are presented and the implications of the vegetational and climatic record are discussed. The traditional view of a smooth progress towards more temperate conditions following the Younger Dryas is not supported; between 10 180 and 9120 B.P., three cooler periods are inferred, the earliest of which may belong to a terminal phase of the Younger Dryas. Comparative pollen ‘influx’ data strongly suggest thatQuercus,Ulmus andAlnus were not present locally. As a working hypothesis, it is suggested that the demise of woodland, from ca. 5950 B.P., was a result of exposure. Pollen indicative of human impact was probably derived from areas of lowland agricultural activity from ca. 5330 B.P. onwards.     

Taxonomic status of the hominid mandible KNM-ER TI 13150 from the middle pliocene of tabarin,in Kenya

A partial mandible with two molars intact was recovered between 1981 and 1984 from deposits of the Middle Pliocene at Tabarin, in Kenya. It has been described and assigned toAustralopithecus cf.afarensis Johanson, White, andCoppens, 1978, with the condition that if ‘A. afarensis’ is revised, then the attribution may change. The taxon ‘A. afarensis’ was found to be invalid and was revised. The smaller specimens of ‘A. afarensis,’ to which the Tabarin mandible was said to be similar, were redescribed asHomo antiquus Ferguson, 1984. Since the Tabarin mandible andH. antiquus are successive transients of the same gens and are allopatric, the Tabarin hominid population is described as an earlier chronosubspecies,Homo antiquus praegens ssp. n.     

Semioticians Make Strange Bedfellows! Or, Once Again: “Is Language a Primary Modelling System?”

Like other sciences, biosemiotics also has its time-honoured archive, consisting of writings by those who have been invented and revered as ancestors of the discipline. One such example is Jakob von Uexküll. As to the people who ‘invented’ him, they are either, to paraphrase a French cliché, ‘agents du cosmopolitisme sémiotique’ like Thomas Sebeok, or de jure and de facto progenitor like Thure von Uexküll. In the archive is the special issue of Semiotica 42. 1 (1982) edited by the late Sebeok and introduced by Thure von Uexküll. It is in the opening essay that Thure von UexküIl tries to restore Jakob von Uexküll’s role as a precursor of semiotics by negotiating the Elder with Saussure and the linguistics-oriented ‘semiology’ in his wake. However, semiotic mapping, in the strictly ‘disciplinary’ sense, of Jakob von Uexküll is no easy task because he ‘knew neither Peirce nor Saussure and did not use their terminology’ (Thure von Uexküll 1982,2). Because Thure prefers to call the Elder’s science ‘general semiotics’ (Thure von Uexküll 1982), this paper begins by assessing Thure von Uexküll’s semiotic configuration of Jakob, probe into the force and limits of the linguistic analogy, revisit the already time-honoured debate on the primary and secondary modelling systems, which was made famous by the Moscow-Tartu semioticians in the early 1970s, but severely criticized by Sebeok and his followers. The paper engages Sebeok from several fronts, directed first at his relegation of the Saussurian linguistic model, then at his critique of the Primary Modelling System, and finally at his reservation about evolutionism in light of the current debate on gene/meme co-evolution. Paper presented at the Eighth Annual International Gatherings in Biosemiotics University of the Aegean, Syros, Greece, 23–28 June 2008     

T’ef (<Emphasis Type="Italic">Eragrostis tef</Emphasis>) in Ancient Agricultural Systems of Highland Ethiopia

T’ef ( Eragrostis tef ) in Ancient Agricultural Systems of Highland Ethiopia. T’ef (Eragrostis tef) has been cultivated in the Horn of Africa for at least 2,000 years. The earliest known agricultural systems in this region date to the Pre-Aksumite period (800–400 b.c.) and appear to have focused on Near Eastern crops, with indigenous African species increasing in importance during Aksumite times (400 b.c.–a.d. 700). While palaeoethnobotanical data are available from Pre-Aksumite and late Aksumite periods, macroscopic botanical remains from the site of Ona Nagast, northern Ethiopia, provide a first glimpse of agricultural systems dating to Proto-Aksumite (400–50 b.c.), Early to Classic (50 b.c.–a.d. 340), and Post-Aksumite (a.d. 700–900) times. Archaeological t’ef remains from Ona Nagast are examined in detail. Guidelines are developed for the identification of t’ef grains preserved on archaeological sites, with a focus on how to differentiate them from seeds of wild Eragrostis species. Charring experiments reveal that in some cases t’ef may not survive high temperatures tolerated by larger cereal grains, such as wheat and barley. The domestication history of t’ef appears to be different from some other cereals, a factor which may explain the preponderance of indeterminate Eragrostis seeds in archaeological samples. Selection of large seed size and intensified tillage were not key factors in t’ef domestication. Early cultivators were likely selecting for increased branching and higher percentage seed set under conditions of minimal tillage.     

A dynamic model of motor basal ganglia functions

 Fast aiming movements were measured in a choice reaction paradigm in a healthy control group and in Parkinsonian patients. The patients were tested without (‘off ’) and with 3,4-dihydroxyphenylalanine (‘on’) (L-dopa) medication. The movement trajectories were used to estimate the parameters of a dynamic linear model. The model is based on the functional structure of the basal ganglia-thalamocortical circuit with direct and indirect pathways linking the putamen to the basal ganglia output nuclei (Albin et al. 1989). The output of the circuit is connected to a model for the motor neuron-musculo-skeletal system. The gain k _d for the direct pathway and the gain k _i for the indirect pathway were estimated. They were found to be significantly decreased for Parkinsonian patients in ‘off ’ compared with the control group. L-dopa therapy in Parkinsonian patients increased the gains of the direct and the indirect pathway almost to normal values which implies that the long-term dopamine level in the striatum was excitatory for the direct and for the indirect pathway. This result is restricted to movements of correct size. For movements of diminished size, which are typical for Parkinsonian patients, the model predicts that the dopamine level in the striatum is excitatory for the direct pathway but inhibitory for the indirect pathway. The simulated values for neuronal activities are in agreement with expected values according to the experimental data. The proposed model of the ‘motor’ basal ganglia thalamocortical circuit implies that information about biomechanical properties of the musculo-skeletal system is stored in the ‘motor’ basal ganglia-thalamocortical circuit, and that the basal ganglia are involved in computation of the desired movement amplitude. Received: 24 April 1996/Accepted in revised form: 25 February 1997     

Different behaviour of crop species in phosphate absorption

Summary Isotopic-dilution technique was used to study the differential phosphate absorption abilities of several field crops. The response of crops was found to be inversely related to the ratio of phosphorus absorbed from the soil to that absorbed from the fertilizer. The ‘A’ values ranged from 2.03 ppm for mustard to 21.77 ppm for soybeans. A highly significant correlation (r=0.77**) was found between per cent yields and the ‘A’ values.     

Bitterness inheritance in apricot (<Emphasis Type="Italic">P. armeniaca</Emphasis> L.) seeds

Seed bitterness, due to cyanogenic glucosides, has been reported in apricot as a recessive trait, being determined by a single gene. In this study, 21 F1 and 10 F2 populations from parents with either bitter or non-bitter (‘sweet’) phenotype were tested by seed tasting. Both the ‘bitter’ and the ‘sweet’ phenotypes were represented in populations from ‘bitter × bitter’ and ‘sweet × sweet’ crosses, as well as from self-pollination of either bitter- or sweet-seeded trees, providing evidence that more than one gene is involved in this trait. Ten populations showed segregation ratios inconsistent with a monofactorial inheritance of seed taste with the ‘sweet’ trait dominant over the ‘bitter’. On the other hand, data from spectrophotometric assays indicate that seed cyanoglucoside content cannot be regarded as a quantitative trait. All the observed segregation ratios can be explained by an inheritance mechanism based on five, non-linked genes, involved in two distinct biochemical pathways. Three genes would control different steps in an ‘additive’ pathway (either the biosynthesis of cyanoglucosides, or their transport, or both) leading to accumulation of these metabolites in seeds: homozygosis of recessive alleles of at least one of them would result in the sweet phenotype. Two more genes would provide a cleaving activity, participating to cyanoglucoside catabolism; heterozygosis or homozygosis of dominant alleles at these loci would produce the ‘sweet’ phenotype, while homozygosis for recessive alleles of at least one of them would interrupt the catabolic pathway, leading to the ‘bitter’ trait, if associated with the anabolic function.     

Distinguishing wild and domestic wheat and barley spikelets from early Holocene sites in the Near East

Identifying morphologically domestic cereals is essential to understanding the origins of agriculture. Charred spikelet bases provide the best evidence for distinguishing wild from domestic varieties of emmer, einkorn and barley; however until now, identification criteria have not been agreed upon or well established. We examined more than 20,000 remains of charred spikelets from eleven early Holocene sites in the Near East, classing them into nine groups. We show that damage and fragmentation of wheat spikelets probably due to dehusking makes identification problematic, and only when the abscission scar is well preserved is it possible to distinguish wild spikelets which shatter from domestic spikelets which adhere and separate during threshing. Barley spikelets were found to be less damaged and more easily identified, perhaps because the processing was less damaging. Einkorn was dominant over emmer on early sites, whereas on later sites emmer was dominant. Identifications presented here from eleven sites date from approximately 13000 to 8200 cal b.p. They give an incomplete picture, but no domestic cereals were identified during the PPNA (Pre-pottery Neolithic A). Early PPNB Aswad produced domestic barley but at other sites for this period the evidence is inconclusive. Unequivocal signs of domestic emmer spikelets appear during the Middle PPNB about 10,000 years ago but wild forms continue as part of the crop after this period. These conclusions are based on limited data. Future studies will undoubtedly produce a more accurate picture.     

Genetic variability and structure of natural and domesticated populations of Caribbean pine (Pinus caribaea Morelet)

 Isozyme analysis of seed samples derived from natural and managed populations of the tropical pine Pinus caribaea vars ‘bahamensis’ and ‘caribaea’ was used to assess population genetic structure in its native range and to detect changes occurring during early domestication of the species. Baseline data from natural populations of the two varieties showed that populations sampled as seed are characterized by high gene diversity (mean H_e=0.26) and a low level of inbreeding ( mean F_is=0.15). A UPGMA tree of genetic relatedness among populations indicates that the two varieties represent distinct evolutionary units. Within each variety there is significant differentiation among populations, and this is greater for the more fragmented populations of var ‘bahamensis’ (F_st=0.08) than for var ‘caribaea’ (F_st=0.02). Seed from a seed orchard population of var ‘caribaea’ established within its natural range showed no change in genetic diversity but did show a reduced inbreeding coefficient (F_is=0.09) compared with its progenitor populations, suggesting a decrease in selfing and/or biparental inbreeding. A bulked seed sample from an exotic plantation of var ‘bahamensis’ in Australia displayed a large increase in the inbreeding coefficient (F_is=0.324) compared with that found in natural populations, possibly due to elevated self-fertilization. Finally, a bulked seed sample from an exotic plantation population of var ‘caribaea’ from China showed enhanced genetic diversity, an increase in the inbreeding coefficient and more linkage disequilibrium than its presumed progenitor populations. It was also genetically divergent from them. RFLP analysis of chloroplast DNA variation in the Chinese sample suggested that seeds of the related taxa P. elliottii and P. taeda, or seeds derived from hybridization with these taxa growing in the seed production area, had been included in the seed crop during harvesting. We conclude that monitoring of appropriate genetic markers may be an effective means of identifying potentially deleterious genetic changes occurring during forest tree domestication. Received: 10 August 1998 / Accepted: 8 September 1998     

Molecular diversity at 18 loci in 321 wild and 92 domesticate lines reveal no reduction of nucleotide diversity during Triticum monococcum (Einkorn) domestication: implications for the origin of agriculture

The diploid wheat Triticum monococcum L. (einkorn) was among the first crops domesticated by humans in the Fertile Crescent 10,000 years ago. During the last 5,000 years, it was replaced by tetraploid and hexaploid wheats and largely forgotten by modern breeders. Einkorn germplasm is thus devoid of breeding bottlenecks and has therefore preserved in unfiltered form the full spectrum of genetic variation that was present during its domestication. We investigated haplotype variation among >12 million nucleotides sequenced at 18 loci across 321 wild and 92 domesticate T. monococcum lines. In contrast to previous studies of cereal domestication, we sampled hundreds of wild lines, rather than a few dozen. Unexpectedly, our broad sample of wild lines reveals that wild einkorn underwent a process of natural genetic differentiation, most likely an incipient speciation, prior to domestication. That natural differentiation was previously overlooked within wild einkorn, but it bears heavily upon inferences concerning the domestication process because it brought forth 3 genetically, and to some extent morphologically, distinct wild einkorn races that we designate here as alpha, beta, and gamma. Only one of those natural races, beta, was exploited by humans for domestication. Nucleotide diversity and haplotype diversity in domesticate einkorn is higher than in its wild sister group, the einkorn beta race, indicating that einkorn underwent no reduction of diversity during domestication. This is in contrast to findings from previous studies of domestication history among more intensely bred crop species. Taken together with archaeological findings from the Fertile Crescent, the data indicate that a specific wild einkorn race that arose without human intervention was subjected to multiple independent domestication events.     

Vegetation history,climate and human impact over the last 15,000 years at Lago dell’Accesa (Tuscany,Central Italy)

Interdisciplinary studies of the sediments of Lago dell’Accesa started in 2001. We present here results from the palynological study. The pollen diagram provides a record of vegetation and climatic change spanning over 15,000 years. The oldest pollen spectra show a late-glacial steppe vegetation typical of central and southern Italy during this period. The Late-glacial Interstadial, interrupted by two cooling events, is dominated by open deciduous oak forests. The Younger Dryas is represented by 150 cm of sediment and shows the presence of steppic vegetation. The Holocene vegetation is characterised by alternating dominance of deciduous oaks and Quercus ilex. The three zones characterised by Q. ilex are accompanied by peat layers marking lake-level lowering at ca. 8600–7900, 4600–4300 and 3700–2800 cal b.p. Between approximately 9000 and 6000 cal b.p. extensive Abies-forests existed on the Colline Metallifere located 15–20 km to the north and northeast of the lake. Local fir populations may also have existed by the lake. Human impact starts at approximately 8000 cal b.p. during the Neolithic period, and increases at ca. 4300 cal b.p. Castanea and Juglans pollen is recorded from ca. 2800 cal b.p. The impact of the Etruscan settlement near the lakeshore is shown in the increasing values of arable crops, species of secondary forest canopy (Ericaceae, Pinus, Pistacia, Myrtus) and anthropogenic indicators (Chenopodiaceae, Plantago lanceolata, Rumex etc).