Phenological behaviour of <Emphasis Type="Italic">Quercus</Emphasis> in Ourense (NW Spain) and its relationship with the atmospheric pollen season

Knowing the beginning of the pollen season is of particular importance to people allergic to a given pollen. Phenological studies in combination with aerobiological studies enable us to observe the relationship between the reproductive phenology of the species and their airborne pollen curves. They also enable us to study the relationship between the release of pollen from anthers and its presence in the atmosphere. The objectives of this study are (1) to determine chill and heat requirements for different phenological phases in the Quercus species present in Ourense (Spain), (2) to verify the synchronization between the flowering event and the presence of Quercus pollen in the air, (3) to select the best criterion for determining the onset of the pollen season according to phenological data, and finally, (4) to obtain models enabling us to determine the onset of the pollen season. Pollen monitoring was carried out from 1993 to 2001, by means of a 7-day Lanzoni VPPS pollen trap and, in order to characterize the floral phenophase of different Quercus species, a sampling method was applied from February to June 2001. We observed that climatic and biotic factors can produce a delay in the onset of the pollen season. Temperatures above 8 degrees C and between 6 degrees C to 8 degrees C are good threshold temperatures for estimating chill and heat requirements respectively. Medium-distance transportation of Quercus robur and Quercus pyrenaica pollen from areas 10-30 km away influences the concentrations of Quercus pollen registered in Ourense.     

Differences in atmospheric trees pollen seasons in winter, spring and summer in two European geographic areas, Spain and Italy

The global climate change reported over recent years may prompt changes in the atmospheric pollen season (APS). The aim of this study is to evaluate the possible impact provoked by meteorological conditions variations at different seasons of the year or different geographical areas on APS. Alnus, Betula and Castanea atmospheric pollen seasons and trends during the last 17 years at Ourense and Vigo (Galicia—NW Spain) and Perugia (Italy) were analysed. Possible incidence of the meteorological trends observed in the different cities on the atmospheric pollen seasons and the chill and heat requirements were evaluated. Pollen data from Ourense, Vigo and Perugia (1995–2011) were used. Pollen sampling was performed using LANZONI VPPS 2000 volumetric traps (Hirst in Ann Appl Biol 36:257–265, 1952), placed on top of different buildings at a similar height from the ground. Several methods, dates and threshold temperatures for determining the chill and heat requirements needed to trigger flowering were tested. Different temporary order in the pollination sequence was observed between the three pollen types studied in the three sites. Alnus flowers few days in advance in Ourense respecting to Vigo and 1 month earlier than Perugia. The Betula flowering start date in Ourense and Vigo is almost simultaneous, taking place only 5 days in advance with respect to Perugia. Finally, scarce differences in the APS onset of Castanea were detected between the three cities. The variations observed among the two areas (Umbria, Italy and Galicia, Spain) in the onset of pollen season in the winter or spring flowering trees could be explained by differences in the thermal requirements needed for flowering as consequence of the climatic conditions recorded during the previous period to flowering. The length of the chilling and heat period as well as the thermal requirements obtained showed differences between geographical areas. The chill requirements accumulated were higher in Perugia than Ourense and Vigo. By contrary, the lowest heat accumulation was achieved in Perugia. The observed trends in the APS characteristics and the weather-related parameters were not homogeneous both in the pollen types and sites. The pollen index of Betula and Castanea pollen in Ourense shows a significant trend to increase.     

Use of <Emphasis Type="Italic">Quercus ilex</Emphasis> subsp. <Emphasis Type="Italic">ballota</Emphasis> phenological and pollen-production data for interpreting <Emphasis Type="Italic">Quercus</Emphasis> pollen curves

Although aerobiological data are often used in phenological research as an indicator of flowering, airborne pollen concentrations are influenced by a number of factors that could affect pollen curves. This paper reports on a study of various aspects of reproductive biology in Q. ilex subsp. ballota, together with environmental factors influencing pollen release and transport, with a view of achieving reliable interpretation of Quercus pollen curves in Ourense (NW Spain). Aerobiological data were recorded from 2002 to 2004 at two sites in the province of Ourense. From 1st February to the end of the flowering period, phenological observations were carried out on 19 trees from the Q. ilex subsp. ballota population found in the Ourense area. Pollen production was calculated for the same trees. The chilling and heating requirements for triggering development were also calculated. The mean flowering period lasted 11-15 days. Reduced pollen output per catkin and, especially, a reduced number of catkins per tree in 2003 and 2004, prompted a marked decline in overall pollen production. Major differences observed in Q. ilex subsp. ballota pollen curves were attributed to the considerable influence both of weather conditions during pollination and pollen production. In years with high pollen production and weather conditions favouring pollen release, Q. ilex subsp. ballota contributed almost 10% to the total Quercus pollen curve. Around 20% of the pollen trapped was captured before or after flowering periods.     

<Emphasis Type="Italic">Betula</Emphasis> pollen: One of the most important aeroallergens in Ourense,Spain. Aerobiological studies from 1993 to 2000

This study presents the results obtained for airborne Betula pollen between 1992 and 2000 in Ourense, Spain, sampled by volumetric spore-trap (LANZONI VPPS2000). Annual and year-on-year variations were analyzed, and a statistical study of the correlation between daily counts and several meteorological parameters was performed. Birch pollen is present in the atmosphere during March and April in Ourense. Significant differences were observed among the different years. Values obtained for the correlation coefficient between Betula pollen counts and the various meteorological parameters studied indicate, for Ourense, a positive correlation between pollen count and both temperature and sunlight. A negative correlation was recorded for relative humidity. Temperature is thus the determining factor for flowering onset and intensity. Regression equations included values for the days prior to pollen concentration measurement in order to optimize results.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

Temperature requirements of Alnus pollen in Spain and Italy (1994-1998)

Woody plants in temperate regions, in order to prevent the water in their cells from freezing, interrupt their growth entering into a physiological state called "dormancy". Trees also have a heat requirement that must be fullfilled before actual growth is resumed and pollination occurs. The objective of this study is to ascertain the influence that a climatic parameter such as temperature exerts on Alnus, in order to identify the start of the dormancy period, its duration and the consequent heat requirement that triggers flowering in three European regions located at the same latitude. In this regard, we chose two areas in Spain (Vigo and Santiago de Compostela) and one in Italy (Perugia), since they have different temperature patterns, ranging from mild in the case of Vigo to cold in that of Perugia, including an intermediate temperature pattern (Santiago). Monitoring in the three stations was carried out by means of a 7-day Lanzoni VPPS 2000 pollen trap. Alnus began its pollen season between the second and third week of January in Spain, while in Italy it began during the first week of February. During the four years under study, in order to overcome the dormancy period, Perugia needed an average of 962 chilling hours (CH), Santiago 622 CH, and Vigo had the smallest chilling accumulation with only 460 CH. In the case of heat accumulation, we found that Perugia had the smallest requirement and Vigo the largest, thus Perugia needed an average of 42 (growth degree days) GDD, Santiago around 50 GDD and Vigo 62 GDD.     

Reproductive biology of Boswellia serrata, the source of salai guggul, an important gum-resin

Detailed studies were carried out on the phenology, floral biology, pollination ecology and breeding system of Boswellia serrata Roxb. (Burseraceae) the source of 'salai guggul'. The trees remain leafless during the entire period of flowering and fruiting. The inflorescence is a terminal raceme and produces up to 90 bisexual, actinomorphic flowers. On average a flower produces 10 044 ± 1259 starch-filled pollen grains. About 85% of the fresh pollen grains are viable; the pollen to ovule ratio is 3348 : 1. The stigma is of the wet papillate type. The style is hollow with three flattened stylar canals filled with a secretion product. The stylar canals are bordered by a layer of glandular canal cells. The inner tangential wall of the canal cells shows cellulose thickenings. The ovary is trilocular and bears three ovules, one in each locule. Flowers offer nectar and pollen as rewards to floral visitors. The giant Asian honey bee ( Apis dorsata ) and A. cerana var. indica (Indian honey bee) are the effective pollinators. The species is self-incompatible and the selfed pollen tubes are inhibited soon after their entry into the stigma. Self-pollen tubes develop a characteristic 'isthmus' as a result of enlargement of the tube soon after emergence through the narrow germ pore. Cross-pollinated flowers allowed normal pollen germination and pollen tube growth, and resulted in fruit- and seed-set. Under open pollination fruit-set was only about 10%. Although manual cross-pollinations increased fruit set, it was only up to about 20%. Low fruit set appears to be the result of inadequate cross-pollination and other constraints, presumably limitation of available nutrients.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 73–82.     

Flowering in a bamboo, Melocanna baccifera (Bambusoideae: Poaceae)

Melocanna baccifera (Roxburgh) Kurtz ex Skeels, a species of bamboo introduced to Sri Lanka from India, flowered and set fruit during 2001–2002. Culms that flowered and set fruit died. The incidence of flowering is significant in that flowering took place close to the predicted mast flowering in 2007. At the onset of flowering, inflorescences were predominantly staminate. But later in 2002, bisexual and pistilate flowers also developed leading to fruiting. Both protandry and protogyny were observed in the bisexual florets. Floral characters indicated that the species was mainly out-crossing. Although anther dehiscence released pollen and stigmas were exerted, pollination of stigmas was inefficient. The few stigmas that were naturally pollinated showed limited pollen tube growth. However, fruit set took place. Fruits were very rarely seen to germinate naturally. Many fruits were devoid of an embryo, indicating that parthenocarpy and/or apomixis may have taken place. Excised embryos germinated in vitro .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 287–291.     

Bauhinia larsenii, a fossil legume from Guangxi, China

Fossil leaves and a branch with a leaf and a pod attached have been discovered from Ningming County, Guangxi Autonomous Region, China, and are described as Bauhinia larsenii sp. nov. (Leguminosae: Cercideae). The strata from which the Bauhinia fossils were collected, namely the Ningming Formation, were dated as Late Eocene–Oligocene based on a combination of pollen, fauna and flora. The affinity of the new species to Bauhinia section Micralvesia subsection Viridescentes species is also discussed.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 437–440.     

Hesperis turkmendaghensis (sect. Hesperis) (Cruciferae/Brassicaceae), a new species from the Central Anatolia region, Turkey

Hesperis turkmendaghensis A.Duran & A.Ocak sp. nov. (Cruciferae) is described and illustrated from Anatolia, Turkey. The species grows under mixed forest, open forest and shady slopes in Türkmen Dağı (B3 Eskişehir) in Central Anatolia. It is closely related to H. matronalis L. ssp. matronalis , an endemic confined to Central Anatolia. Diagnostic morphological characters are discussed. Notes are also presented on its ecology, biogeography and conservation status. In addition, the pollen characteristics and seed coat surface of H. turkmendaghensis and H. matronalis are examined by SEM.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 239–247.     

Reproductive biology in Anagyris foetida L. (Leguminosae), an autumn–winter flowering and ornithophilous Mediterranean shrub

As most plants of the Mediterranean region bloom in spring, there have been few studies of the reproductive biology of species with autumn–winter flowering. In this study, we investigate the breeding system of Anagyris foetida , one of the few shrubs that blooms at this time. The floral, phenological, and reproductive aspects of two populations of this Mediterranean legume from south-west Spain were studied via field and laboratory experiments. The variability of fruit and seeds was studied in another 12 Iberian populations with respect to certain meteorological parameters (temperature and rainfall). Anagyris foetida shows cauliflory, marked floral longevity, and adichogamy. The peak of flowering is in January–February. It is self-compatible, with no clear advantage of cross- over self-pollination, and with virtually no autonomous self-pollination. This is because the stigma, like some other legumes, prevents the germination of pollen if its surface is not ruptured by pollinators. The number of seeds per fruit under natural pollination was positively correlated with the total rainfall during the fruiting period (from January to May), and significantly influenced the percentage of fruit weight represented by the pericarp, in the sense that the smaller the number of viable seeds in the fruit, the greater the percentage of pericarp weight.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 519–532.     

On the presence of extracellular calcium oxalate crystals on the inflorescences of Araceae

This study presents a survey of the species of the Araceae where extracellular production of calcium oxalate crystals has been observed and discusses the patterns of production of the crystals in different genera. For all Araceae studied using SEM, the oxalate crystals exuding on the epidermal surface correspond to extended aggregate/druses or crystal sand and the oxalate crystals mixed with pollen correspond to raphides or styloids (prismatic crystals). The type of crystals associated with pollen varies among genera. However, the presence of crystals associated with pollen is a specific rather than a generic characteristic. Our results show that the presence of raphides mixed with pollen seems to be a widespread phenomenon in the aroid family.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 181–190.     

A study of Quercus pollen in the Derby area, UK

The major allergenic pollen prevalent in the Derby air in May is Quercus pollen which has been monitored volumetrically from 1970–1997. Quercus pollen levels in Derby are increasing, showing an established long term trend, with 1995 being an exceptionally high year. There is now an earlier start date and a longer seasonal duration. The mean Quercus diurnal periodicity for 1991–1997 shows a peak at 15.00 hours.A detailed study of the 1990–1997 seasons established that a maximum temperature of 20 °C or above, at the usual time of flowering, occasions the start of the Quercus pollen season. Average May temperature and drought in the previous June and July are important factors in determining Quercus pollen totals. Predictions for the forthcoming seasons were produced which compared favourably with the actual pollen totals.     

The effectiveness of some mechanisms of reproductive isolation in Arum maculatum and A. italicum (Araceae)

Mechanisms of reproductive isolation were studied in two species of Arum , A. italicum and A. maculatum , growing in England and the south of France. The study focused on three potential mechanisms for reproductive isolation: the effectiveness of dichogamy as a barrier to autogamy; the ability of self and outcrossed pollen to germinate on stigmas at different stages of anthesis and to effect pollination; and postzygotic barriers to selfing. Dichogamy was found to provide a very effective barrier to within-inflorescence selfing in these species, as no seeds were produced by spontaneous self-pollination (i.e. autogamy) in any population of either species. However, the study found that geitonogamy (cross-pollination between inflorescences of a same individual or clone) was possible, as genotypes frequently produced several inflorescences and stigmas were found to be receptive to pollen from before anthesis until their contraction at the end of the female phase of flowering. Hand pollination with self pollen from clone inflorescences produced as numerous and heavy seeds as outcross pollination. In addition, the germination and growth of geitonogamously produced seed was similar to that of outcrossed seed, suggesting that this potential postzygotic barrier is absent or weak. These findings suggest that geitonogamous seed production may be unrestricted by significant pre- or postzygotic barriers. The possible advantages of geitonogamy in Arum are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 323–328.     

Reproductive strategies of Strobilanthes kunthianus, an endemic, semelparous species in southern Western Ghats, India

Strobilanthes kunthianus (Acanthaceae) is a semelparous species with synchronized flowering and mast seeding once in 12 years. As semelparous plants have only one chance to reproduce, they are expected to develop effective strategies to prevent reproductive failure. The reproductive strategies of S. kunthianus were investigated by studying the floral traits, pollination biology, and breeding system that are critical for reproductive success. The species exhibits a series of floral traits: (1) gregarious flowering attracts a large number of Apis cerana indica , the major pollinator; (2) the stigma is sensitive to touch by the pollinator; in fresh flowers, the receptive surface faces the entry path of the incoming bee, facilitating pollen deposition; as an immediate response, the stigma curves backwards moving the receptive surface away from the path of the exiting bee, thus preventing autogamy and interference in pollen transfer; (3) flowers remain fresh for 2 days with receptive stigma and nectar and pollen reward. These traits render the species 100% pollination efficient to ensure a high seed set. As the species is self-compatible, the prevailing high degree of geitonogamous pollinations does not interfere with fruit set. The evolution of the adaptive floral traits has facilitated mast seeding in the species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 155–163.     

Little genetic differentiation across Europe between early-flowering and late-flowering populations of the rapidly declining orchid Neotinea ustulata

Amplified fragment length polymorphism (AFLP) was used to characterize genetic diversity of the endangered Burnt Orchid, Neotinea (formerly Orchis ) ustulata . Fingerprinting of Estonian and British populations revealed surprisingly little genetic differentiation between populations but larger amounts of diversity within populations, especially in Britain. The resulting mean F _st value of 0.51 is unusually high for an orchid species. Much of the variation follows a west–east cline across Europe, whereas the much-discussed early- and late-flowering taxa of N. ustulata are considered insufficiently distinct to be viewed as separate subspecies. The later flowering N. ustulata var. aestivalis probably evolved independently on two or three occasions, each time diverging from the earlier flowering nominate race. The identity of the genes underpinning phenology in the species, and the potential selective advantages of phenological divergence, merit further study. Overall genetic diversity within populations is sufficiently high to render impoverishment an unlikely cause of their recent, precipitous decline.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 13–25.     

Influence of precipitation and temperature on airborne pollen concentration in Santiago de Compostela (Spain)

This paper presents data of airborne pollen concentration of Pinus , Platanus , Quercus , Betula , Poaceae and Castanea , the most abundant pollen types, during the last eight years, in Santiago de Compostela (NW Iberian Peninsula). We determined their respective principal pollen release periods and analysed statistically the influence of the main meteorological parameters on their pollen concentration, obtaining a negative correlation with precipitation and a positive correlation with temperature, in the majority of the cases, during these years. A detailed study of accumulated pollen values, in relation to accumulated average temperatures and the precipitation distribution, enabled us to establish a relationship between these parameters and the principal pollination period of each of these taxa.     

Genetic structure of the endangered perennial plant Eryngium alpinum (Apiaceae) in an alpine valley

We investigated the genetic structure of Eryngium alpinum (Apiaceae) in an Alpine valley where the plant occurs in patches of various sizes. In a conservation perspective, our goal was to determine whether the valley consists of one or several genetic units. Habitat fragmentation and previous observations of restricted pollen/seed dispersal suggested pronounced genetic structure, but gene dispersal often follows a leptokurtic distribution, which may lead to weak genetic structure. We used nine microsatellite loci and two nested sampling designs (50 × 50 m grid throughout the valley and 2 × 2 m grid in two 50 × 10 m quadrats). Within the overall valley, F -statistics and Bayesian approaches indicated high genetic homogeneity. This result might be explained by: (1) underestimation of long-distance pollen/seed dispersal by in situ experiments and (2) too recent fragmentation events to build up genetic structure. Spatial autocorrelation revealed isolation by distance on the overall valley but this pattern was much more pronounced in the 50 × 10 m quadrats sampled with a 2-m mesh. This was probably associated with limited primary seed dispersal, leading to the spatial clustering of half-sibs around maternal plants. We emphasize the interest of nested sampling designs and of combining several statistical tools.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 667–677.     

Cannabis sativa: volatile compounds from pollen and entire male and female plants of two variants, Northern Lights and Hawaian Indica

Sixty-eight compounds were identified by coupled gas chromatography and mass spectrometry (GC-MS) in the chemosphere of Cannabis sativa L. pollen and entire male and female plants of two cultivated varieties, Northern Lights and Hawaian Indica. Twenty-one and 28 substances, respectively, were present in pollen of the two forms. To conserve the natural composition of volatiles a delicate headspace method was employed. The two varieties represent different chemotypes which distinguish themselves, in the main quantitatively, in the setup of volatiles from pollen and entire male and female plants. Twenty compounds were monoterpenes, including the five major components: β-myrcene (E)-β-ocimene, terpinolene, β-pinene and limonene; 25 were sesquiterpenes, and the other 23 were of mixed biogenetic origin, including 3-methyl-1-butanol and benzylalcohol which occurred only in pollen; two pyrazines occurred only in Northern Lights females. Besides being of interest in natural products chemistry, the results should have relevance for plant systematics and for the pharmaceutical and technical applications of Cannabis . We demonstrate that the pollen has a distinct chemical character in possessing two exclusive volatiles, while lacking seven compounds occurring in males and females of both variants.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 387–397.     

Pollen fates and the limits on male reproductive success in an orchid population

Male reproductive success in higher plants depends largely on the fates of pollen, but current methodologies have given only partial insights into this important aspect of plant mating. We present a detailed analysis of the proportions and absolute amounts of stained pollen involved in six key fates for the hawkmoth-pollinated African orchid Disa cooperi . Despite being packaged into pollinaria, high proportions (> 0.95) of the pollen removed from anthers were lost during transport by hawkmoths in both years. The proportion of pollen lost correlated positively with the number of pollinaria removed from a plant, so that pollen export did not vary with pollen removal. Most pollen was dispersed to neighbouring plants, with rare long-distance dispersal up to 65 m. Of the pollen that reached stigmas during both years, roughly equal amounts were involved in facilitated self-pollination vs. cross-pollination, but the relative proportions of these fates differed between years. Contrary to expectation, we found that self-pollination between flowers did not increase with the number of open flowers, even though moths probed significantly more flowers on larger plants. However, during both years the fraction of removed pollen exported to other plants declined significantly with increasing self-pollination on the source plant, indicating that once self-pollination occurred it reduced (discounted) subsequent pollen export opportunities. The packaging of pollen into pollinaria in orchids appears to increase overall transfer efficiency by at least an order of magnitude relative to plants with granular pollen. Nevertheless, considerable uncertainties remain in the male reproductive success of individual orchids. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 175–190.