The system of grasses (Poaceae) and their evolution

Originally published in Russian as number 37 of Komarov Readings, 1987. Translated by Victoria V. Michaelova, Brigham Young University, Provo, Utah 84602. Translation edited by Arthur Cronquist, New York Botanical Garden, New York, and approved by the author. A few additions have been made to account for very recent literature.     

The effects of large-scale atmospheric fields on the cypress pollen season in Tuscany (Central Italy)

This research was performed for the purpose of analysing the relationships between large-scale meteorological information, in particular the North Atlantic Oscillation (NAO) index and the Sea Surface Temperature (SST), and the timing and magnitude of the Cupressaceae pollen season in the Pistoia district of Central Italy. The results demonstrated that in specific periods of the year, the NAO index, by partially determining the distribution of the main meteorological variables over the study area, is negatively correlated with the start and the end, as well as the peak day of pollen concentration. Pollen data were also correlated with the SST of the North Atlantic Ocean east of the Azores for the September–December period of the previous year, which is significant for exploring possibilities in terms of predicting the timing and magnitude of the cypress pollen season. The analysis of such meteorological variables and indices could be used to improve the existing forecasting systems of the phenology of the cypress pollen season. Moreover, the possibility of using meteorological information freely available on internet could cut costs and reduce spatial and temporal representativeness limitations relating to weather monitoring in loco.     

Attribute correlations in the Poaceae (grasses)

Investigation of correlations between a number of two-state attributes for a sample of grass genera revealed that a high proportion of the correlations were statistically significant. There were proportionately more correlations between attributes from the same organ than between attributes from different organS. Furthermore, attributes associated with the fruit and leaf were found to be more closely correlated with attributes other than those of the flower and spikelet, thereby indicating fruit and leaf attributes to be important in the classification of the grasseS. It is considered that the correlations have arisen in part as a result of the phylogenetic origins of the genera and in part as a response to natural selection determining distribution of genera adapted to present-day habitats.     

Airborne pine (Pinus spp.) pollen in the atmosphere of Perugia (Central Italy): Behaviour of pollination in the two last decades

The phenology of many species, which grow intemperate climate, is principally regulated bythe temperature and the plants respond withvariations in the beginning, in the durationand in the intensity of the various phenophasestowards every climate change. We have analysedthe data of Pinus pollination in Perugia,Central Italy, during last 2 decades(1982–2001), in a period during which theannual mean temperature significantly increasedby about 0.8 °C.The pine pollination started, on average,between the end of March and mid-April andended in the last days of June, with a meanduration of 65 days. The start dates showed asignificant negative correlation with theaverage air temperature in March andsignificant trends towards an earlier beginningof pollination by 18 days (–0.9 day/year) and ashorter duration of the pollen season by 10days (–0.6 day/year) were found over thestudied period. Moreover, the trend of thedaily pollen counts showed, on average, analmost normal distribution, but the analysis ofeach yearly trend revealed significantdifferences correlated with the meantemperature during the pollen season. Theseobserved trends in pine pollination suggest theuse of aerobiological monitoring of thisairborne pollen as indicator of temperaturechange in Central Italy over a relatively longperiod.     

The results of 2-year pollen monitoring of an urban network in Perugia,Central Italy

Continuous pollen monitoring of an urban network consisting of three stations has been undertaken for a period of 2 years in Perugia, central Italy. The aim has been to establish whether the Perugia pollen trap, active since 1983, is still representative of the area following recent urbanisation. Quantitative differences were found between the stations, reflecting different vegetational areas, but only slight differences were detected in relation to the timing of the principal period of pollination. Therefore, although individual pollen traps are necessary to characterize fully the different areas, one trap is sufficient to determine the key allergenic thresholds in the studied area.     

The results of 2-year pollen monitoring of an urban network in Perugia,Central Italy

Continuous pollen monitoring of an urban network consisting of three stations has been undertaken for a period of 2 years in Perugia, central Italy. The aim has been to establish whether the Perugia pollen trap, active since 1983, is still representative of the area following recent urbanisation. Quantitative differences were found between the stations, reflecting different vegetational areas, but only slight differences were detected in relation to the timing of the principal period of pollination. Therefore, although individual pollen traps are necessary to characterize fully the different areas, one trap is sufficient to determine the key allergenic thresholds in the studied area.     

Correlation between large-scale atmospheric fields and the olive pollen season in Central Italy

Olives are one of the largest crops in the Mediterranean and in central and southern Italy. This work investigates the correlation of the Olea europaea L. pollen season in Perugia, the capital city of the region of Umbria in central Italy, with atmospheric parameters. The aim of the study is twofold. First, we study the correlation between the pollen season and the surface air temperature of the spring and late spring in Perugia. Second, the correlation between the pollen season and large-scale atmospheric patterns is investigated. The average surface temperature in the spring and late spring has a clear impact on the pollen season in Perugia. Years with higher average temperatures have an earlier onset of the pollen season. In particular, a 1°C higher (lower) average surface temperature corresponds to an earlier (later) start of the pollen season of about 1 week. The correlation between the pollen season and large-scale atmospheric patterns of sea level pressure and 500-hPa geopotential height shows that the cyclonic activity in the Mediterranean is unequivocally tied to the pollen season in Perugia. A larger than average cyclonic activity in the Mediterranean Basin corresponds to a later than average pollen season. Larger than average cyclonic activity in Northern Europe and Siberia corresponds to an earlier than average pollen season. A possible explanation of this correlation, that needs further investigation to be proven, is given. These results can have a practical application by using the seasonal forecast of atmospheric general circulation models.     

Phenological and aeropalynological survey in an olive orchard in Umbria (Central Italy)

The olive tree Olea europaea L. is a resource of notable value from an economic and landscape point of view in Central Italy. This study examined two cultivars (Giarraffa and Ascolana) grown in an associative stand in an experimental olive orchard, to verify the possibility of introducing new olive cultivars into Umbria. The analyses were carried out at three different altitudes in the olive orchard following criteria used for non-homogeneous samplings in order to minimize the effect due to steepness. Since the number of trees of the two cultivars is different, the sample sizes were different for the two cultivars. Phenological observations of the above mentioned cultivars and the evaluation of the pollen flow, done by using a portable pollen trap inside the olive orchard, were carried out over two seasons (1998-1999). The start of initial phenological phases differed in the two cultivars, but the differences became less evident during phases of anthesis. In aeropalynological respect, it appears that, in a hill species such as the olive tree, the flowering is correlated with altitude. Comparing the aeropalynological data from the distant pollen trap and those collected in the olive orchard, it is clear that the periods of pollen release are very similar in the two cultivars, and there is a clear synchronism in the peak pollen release. It is concluded that in Central Italy the pollen grains of Giarraffa cultivar are well compatible with the Ascolana cultivar from a phenological and aeropalynological point of view.     

Poaceae pollen in Galicia (N.W. Spain): characterisation and recent trends in atmospheric pollen season

Airborne Poaceae pollen counts are greatly influenced by weather-related parameters, but may also be governed by other factors. Poaceae pollen is responsible for most allergic reactions in the pollen-sensitive population of Galicia (Spain), and it is therefore essential to determine the risk posed by airborne pollen counts. The global climate change recorded over recent years may prompt changes in the atmospheric pollen season (APS). This survey used airborne Poaceae pollen data recorded for four Galician cities since 1993, in order to characterise the APS and note any trends in its onset, length and severity. Pollen sampling was performed using Hirst-type volumetric traps; data were subjected to Spearman’s correlation test and regression models, in order to detect possible correlations between different parameters and trends. The APS was calculated using ten different methods, in order to assess the influence of each on survey results. Finally, trends detected for the major weather-related parameters influencing pollen counts over the study period were compared with those recorded over the last 30 years. All four cities displayed a trend towards lower annual total Poaceae pollen counts, lower peak values and a smaller number of days on which counts exceeded 30, 50 and 100 pollen grains/m³. Moreover, the survey noted a trend towards delayed onset and shorter duration of the APS, although differences were observed depending on the criteria used to define the first and the last day of the APS.     

Universal primers for the amplification of chloroplast microsatellites in grasses (Poaceae)

Attempts to design truly universal primers to amplify chloroplast microsatellites have met with limited success due to nonconservation of repeat loci across widely divergent taxa. We have used the complete chloroplast genome sequences of rice, maize and wheat to design five pairs of primers that amplify homologous mononucleotide repeats across the Poaceae (grasses). Sequencing confirmed conservation of repeat motifs across subfamilies and a preliminary study in Anthoxanthum odoratum revealed polymorphism at two loci with a haplotype diversity value of 0.495. These primers provide a valuable tool to study cytoplasmic diversity in this extensively studied and economically important range of taxa.     

The main pollen season of Parietaria in Genoa (Italy): forecast possibilities

The authors propose a statistical model for evaluating the Beginning of the Main Pollen Season (BMPS) of Urticaceae (cf. Parietaria) in Genoa (Italy). For eight consecutive years (1981–1988) the mean daily concentrations of this pollen grain were studied in relation to the two meteorological factors which are most significant in triggering the beginning of maximal pollination, namely, mean daily air temperature and rainfall. From a multiple-regression statistical analysis, a formula has been extrapolated, which, one the basis of these meteorological data for the month of January, can be used to predict the onset of the BMPS to within nine days.     

The {beta}-amylase genes of grasses and a phylogenetic analysis of the Triticeae (Poaceae)

There are two forms of β-amylase in the Triticeae crop plants wheat, barley, and rye: an endosperm-specific form encoded by two or three closely linked genes, and a tissue-ubiquitous form encoded by a single gene. Both rice and corn have one ubiquitously expressed form encoded by a single gene. This study focuses on two phylogenetic analyses of β-amylase gene sequences. First, a phylogenetic analysis of coding sequences from wheat, barley, rye, rice, and corn was expected to clarify the relationship between the endosperm-specific and tissue-ubiquitous forms of the protein. Instead, it illustrates possible effects of distant outgroups, based on conflicting patterns of character state variation consistent with different root positions. Next, a broad sample of the monogenomic Triticeae was included in a phylogenetic analysis based on sequences from a portion of the tissue-ubiquitous β-amylase gene. The results were compared to existing Triticeae gene trees, among which extensive conflict had been noted in the past. One additional gene tree has not completely clarified the complexity of the group, but has shed additional light on reticulate phylogenetic patterns within the tribe, including relationships involving Eremopyrum, Thinopyrum, and the Triticum/Aegilops group.     

Slipped-strand mispairing in a plastid gene: rpoC2 in grasses (Poaceae)

An exception to the generally conservative nature of plastid gene evolution is the gene coding for the beta" subunit of RNA polymerase, rpoC2. Previous work by others has shown that maize and rice have an insertion in the coding region of rpoC2, relative to spinach and tobacco. To assess the distribution of this extra coding sequence, we surveyed a broad phylogenetic sample comprising 55 species from 17 angiosperm families by using Southern hybridization. The extra coding sequence is restricted to the grasses (Poaceae). DNA sequence analysis of 11 species from all five subfamilies within the grass family demonstrates that the extra sequence in the coding region of rpoC2 is a repetitive array that exhibits more than a twofold increase in nucleotide substitution, as well as a large number of insertion/deletion events, relative to the adjacent flanking sequences. The structure of the array suggests that slipped-strand mispairing causes the repeated motifs and adds to the mechanisms through which the coding sequence of plastid genes are known to evolve. Phylogenetic analyses based on the sequence data from grass species support several relationships previously suggested by morphological work, but they are ambiguous about broad relationships within the family.      

Phylogenetic signals in the realized climate niches of Chinese grasses (Poaceae)

Explaining relationships between species richness and biogeographical patterns over a broad geographic scale is a central issue of biogeography and macroecology. We document the realized climate niches for grasses in China’s nature reserves and discuss its formation mechanism using grass richness data combined with climatic, physiological, and phylogenetic data. Our results suggest that climate niche structure of grasses is phylogenetically conservative for BEP (Bambusoideae, Ehrhartoideae, and Pooideae) and PACMAD (Panicoideae, Arundinoideae, Chloridoideae, Micrairoideae, Aristidoideae, and Danthonioideae) clades along temperature gradients and for Chloridoideae and Panicoideae along precipitation gradients. At the national scale, the divergence patterns of climate niches between two major clades are more distinguishable than between C₃ and C₄ grasses. High rates of climate niche evolution are found in C₄ clades in the subtropical forest region. There appears to be a strong association between elevation gradients and grass diversity: the specific environmental conditions (e.g. energy) and the rapid shifts of climate conditions drive high grass diversification. Evolutionary conservatism of climate niches may be influenced by the specific adaptive ability to changing environmental conditions within NAD-ME/NADP-ME clades. Our results indicate that adaptations to major climate changes may be accomplished by C₄ grass nodes of high climate niche evolutionary rates in China’s nature reserves.     

Evolution of Ac and Dsl elements in select grasses (Poaceae)

We present data on evolution of the Ac/Ds family of transposable elements in select grasses (Poaceae). An Ac-like element was cloned from a DNA library of the grass Pennisetum glaucum (pearl millet) and 2387 bp of it have been sequenced. When the pearl millet Ac-like sequence is aligned with the corresponding region of the maize Ac sequence, it is found that all sequences corresponding to intron II in maize Ac are absent in pearl millet Ac. Kimura's evolutionary distance between maize and pearl millet Ac sequences is estimated to be 0.429±0.020 nucleotide substitutions per site. This value is not significantly different from the average number of synonymous substitutions for coding regions of the Adh1 gene between maize and pearl millet, which is 0.395±0.051 nucleotide substitutions per site. If we can assume Ac and Adh1 divergence times are equivalent between maize and pearl millet, then the above calculations suggest Ac-like sequences have probably not been strongly constrained by natural selection. The level of DNA sequence divergence between maize and pearl millet Ac sequences, the estimated date when maize and pearl millet diverged (25–40 million years ago), coupled with their reproductive isolation/lack of current genetic exchange, all support the theory that Ac-like sequences have not been recently introduced into pearl millet from maize. Instead, Ac-like sequences were probably present in the progenitor of maize and pearl millet, and have thus existed in the grasses for at least 25 million years. Ac-like sequences may be widely distributed among the grasses. We also present the first 2 Dsl controlling element sequences from teosinte species: Zea luxurians and Zea perennis. A total of 10 Dsl elements had previously been sequenced from maize and a distant maize relative, Tripsacum. When a maximum likelihood network of genetic relationships is constructed for all 12 sequenced Dsl elements, the 2 teosinte Dsl elements are as distant from most maize Dsl elements and from each other, as the maize Dsl elements are from one another. Our new teosinte sequence data support the previous conclusion that Dsl elements have been accumulating mutations independently since maize and Tripsacum diverged. We present a scenario for the origin of Dsl elements.     

Cupressaceae pollen in the atmosphere of Ascoli Piceno (Central Italy) and sensitization of allergic subjects

The aim of this work is to study the incidence of pollinosis in the Health District of Ascoli Piceno, Central Italy (U.S.L.24), this being an underestimated pathology from the clinical point of view and also as a result of the recent introduction of this taxa in the National Aeroallergen Network. Since 1990, 5055 patients of both sexes with respiratory symptomatology of suspected IgE mediated aetiology have been examined in our Centre with the Skin Prick Test (SPT) using allergen panels including Cypress; 171 (3.38%) patients were found to be positive to this allergen. These results show that the subjects with symptoms in the period January–March in most cases have a sensitization toCupressus pollen and new studies will evaluate the possibility of specific immunotherapy.     

Cupressaceae pollen in the atmosphere of Ascoli Piceno (Central Italy) and sensitization of allergic subjects

The aim of this work is to study the incidence of pollinosis in the Health District of Ascoli Piceno, Central Italy (U.S.L.24). this being an underestimated pathology from the clinical point of view and also as a result of the recent introduction of this taxa in the National Aeroallergen Network. Since 1990, 5055 patients of both sexes with respiratory symptomatology of suspected IgE mediated aetiology have been examined in our Centre with the Skin Prick Test (SPT) using allergen panels including Cypress; 171 (3.38%) patients were found to be positive to this allergen. These results show that the subjects with symptoms in the period January–March in most cases have a sensitization toCupressus pollen and new studies will evaluate the possibility of specific immunotherapy.     

Cleaning up the grasses dustbin: systematics of the Arundinoideae subfamily (Poaceae)

Among the 12 subfamilies currently considered in the systematics of Poaceae, the Arundinoideae have long been considered as a dustbin group, with a diversity of forms putatively hiding incertae sedis. Because this subfamily has been poorly investigated using molecular markers for the last two decades, the present study provides the first complete phylogeny of the Arundinoideae based on five plastid DNA loci sequenced for 12 genera, and analysed with and without plastome data from previous studies. The refined Arundinoideae appear to be a robust evolutionary lineage of Poaceae, divided into three tribes with some biogeographical patterns: (1) tribe Arundineae, the most heterogeneous tribe, including Eurasian Arundo, Australian Amphipogon and Monachather, and South African Dregeochloa; (2) tribe Crinipedeae (described here), including Crinipes, Elytrophorus, Styppeiochloa and Pratochloa (described here), with a South and East African distribution; and (3) tribe Molinieae, including Hakonechloa, Molinia and Phragmites, with a Eurasian distribution. Despite reduction in size, this small subfamily conserves a high diversity of morphological forms, with several small but highly differentiated genera. Finally, the molecular dating approach provides an evolutionary framework to understand the diversification of Arundinoideae, refuting Gondwanan vicariance between genera and suggesting capability for long distance dispersal.     

Incidence of sensitisation to the pollens of urticaceae (Parietaria), Poaceae and Oleaceae (Olea europea) and pollen rain in Liguria (Italy)

Summary The authors examined 734 sensitised patients living in four localities in Liguria (Genoa, Savona, Pietra Ligure and Sanremo). The commonest source of sensitisation (62.7%) was Urticaceae (Parietaria), followed by Poaceae (52.5%) andOlea europaea L. (24.0%). A survey of airborne pollens revealed a greater presence of Urticaceae and Poaceae in Genoa and of Oleaceae in Pietra Ligure and Sanremo.