Floral biology of plum (Review article)

In the majority of Hungarian orchards of stone fruits, the planting distance is 6-7 m x 4-5 m. As many of the current varieties are self-incompatible, planting designs are applied to provide for adequate pollinisers. As long as differences in blooming time are small, i.e. 3-5 days at most, overlaps of blooming of the associated varieties are sufficient for fruit set.

In sour cherry, one leading variety, Pándy, is self-incompatible and requires two polliniser varieties at least (Ciganyneggy or some sweet cherry varieties). Pándy is, moreover, cross-incompatible with the varieties Debreceni bőtermő, Kántorjánosi and Újfehértói fürtös being all of them self-fertile as most of new varieties recommended, by the way, for being planted to monovarietal blocks.

Among European plums there are varieties registered as male sterile, self-incompatible, parially self-fertile and self-fertile, respectively. For the purpose of cross pollination, the choice of two varieties, at least, to be associated to any variety belonging to the first three groups, is recommended. The number of rows in blocks planted to self incompatible or male-sterile varieties should not be higher than 2-(4). Inter-incompatibility has been observed within the currently recommended assortment, between the varieties Cacanska najbolja and Stanley, only. Chinese-Japanese plums are scarcely represented in Hungarian plantations. Variation of blooming time in varieties is somewhat more pronounced, i.e. 5-8 days. There is but a weak tendency to self-fertility, thus practically, all varieties are considered as self-incompatible, thus the planting of two-row blocks for each of three varieties, at least, are recommended to be associated.

Self-incompatibility and partially self-fertile apricot varieties are recommended to be combined with two polliniser varieties, at least, each planted to two-row blocks. The varieties Ceglédi óriás, Ligeti óriás, Nagykőrösi óriás and Szegedi Mammut are mutually inter-incompatible. Most of the peach varieties grown in Hungary are self-fertile, thus they are planted to large blocks, each. On sites threatened by late spring frost, it is recommended to plant (monovarietal) blocks of 4-6 rows at most. Cross-pollination may increase fruit set even in self-fertile varieties.

The resumption of development in ecodormant buds in terms of establishing a functional vascular connection between the inflorescence primordia and spur tissues in apple trees was investigated. Differentiation of the xylem elements could be observed first in the pedicel of the flower primordium, in the middle of January. Much later (at the beginning of April) there were mature xylem vessels in the wall of the receptacle and, merely a procambial strand for the ovule primordium which was at this time an undifferentiated protrusion of meristematic cells, only. As for phenological development of buds incubated at a temperature of 20 °C, it was the slowest in buds sampled in January, faster in buds sampled in the middle of February and, buds from the middle of March responded very quickly. The function of temperatures needed both for xylem differentiation and for the flower primordium to achieve maturity is pointed out. The nature of frost damage in vessel elements, as well the relationship between chilling requirement and growth features of apple cultivars will be discussed.

Due to the geographical situation of Hungary the introduction of late ripening apricot varieties holds great interest. In apricot production floral bud development during winter, blooming time, and the fertilisation properties are important factors. These characters were studied in six late ripening Romanian apricot varieties (Callatis, Comandor, Litoral, Selena, Sirena, Sulmona) in Szigetcsép representing the northern site of the lowland growing area. During the mild winter of 1997/98 the dynamics of floral bud development in the Romanian varieties was a few days slower during the whole examination period compared to Gönci magyar kajszi. Their meiotic divisions occurred between 1 and 5 February. Next winter the meiotic division started later at 28 February, due to the cold weather. In these conditions the dynamics of bud development was similar in all the varieties. Averaged over seven years blooming of the Romanian varieties started 1-3 days later than in Gönci magyar kajszi. All the Romanian varieties showed self-fertility to some extent, however, application of other pollen donor sources is necessary for the safety of production, with the exception of Callatis.

The morphology of the stigma has been studied in 50 varieties belonging to 6 stone fruit species. The majority of samples had elliptical stigmata with some exceptions with circular form (Duane, Tuleu gras). The surface of the stigma is papillary, flattened in side view (sweet cherry) or bulging (apricot, peach). The suture of the stigma is clearly visible as a depression and the varieties may differ in this respect.

The size of the stigma depends highly from the season, although the varietal differences are maintained. The dimension of stigmatic surface is characteristic for the species expressed in square millimetres: sweet cherry 0.92 to 2.91; sour cherry 1.64 to 2.48; plum 0.83 to 1.80; oriental plum 0.53 to 1.15; apricot 0.57 to 1.69 mm².

The size and morphology of the stigma changes according to varieties too, and it may used in description and identification of varieties. No correlation has been found between the size of stigma and the fertility relations (self-fertility or self-incompatibility) of the respective varieties.

Size and surface morphology of pollen has been studied in 87 twit varieties of 10 fruit species during the period of 1990-1995. No preceding work of that type came to our knowledge, yet.

The samples comprised a wide variety of cultivars included male sterile, self-incompatible, partially self-fertile stone fruits, diploid and hexaploid plums, diploid and triploid apples.

The large number of species and varieties facilitated the comparison of items within and between the respective species.

It was stated that the size, shape and surface morphology of pollen is genetically determined and those data, combined with other variety characters, are suitable for the classification and distinction of varieties.

In assessment of pollen size and shape, their moisture content is crucial. The major diameter of the swollen pollen as well as the length and width of the dry grains are characteristic to species and/or to variety.

The width and shape changes largely with moisture content. Large grains are proper to quince, apricot, peach and almond, medium sizes are found in apple, sweet cherry, sour cherry, European plum, whereas small size is typical to Japanese plums.

The low number of varieties studied does not allow conclusions concerning differences within pears, quinces and almonds as species. In the rest of species, valid differences have been registered as between varieties.

Within species, as apple and plum, the effect of ploidy (i.e. number of chromosomes) was expressed in the size of their pollen. In stone fruit species, the correlation between size. of anthers and size of pollen grains was positive.

Genetic relations between the self-fertile sour cherry varieties of the Pándy type (Debreceni bőtermő, Kántorjánosi, Újfehértói fürtös) as well as the self-incompatible apricots of "giant" fruit size are supposed to be analysed by pollen studies but there did not turn out any decisive conclusion, yet. Other characters also should be considered.

The assembly of pollen characters is decisive in the determination of the variety. The ratio of empty pollen grains, the grain size and the density as well as the size of the pits on the surface are best suited to distinguish pollen lots.

Intensive research has been implemented on different aspects of the bee pollination of pear in Hungary in the past decade, extending to the following topics: the effect of the opening sequence of the flowers within the inflorescence of pear cultivars on the effectiveness of bee pollination; the nectar production of flowers of pear cultivars; intensity of honeybee visitation and their foraging behaviour at the flowers of pear cultivars; flower constancy of honeybees to pear plantations; competition by weeds in bloom for bee pollination in flowering pear orchards; and some aspects to the bee pollination strategies of pear plantations (the effect of the limitation of bee pollination period on the set and the yield, possible compensation of frost damage of flowers by bee pollination in pear orchards, the optimal size of single cultivar block in pear plantations). The results of these studies are reviewed in this paper and some important aspects that need further studies are outlined.

Detailed studies were made on the nectar production of 44, 16 and 18 pear cultivars, respectively, in a cultivar collection of pear during three consecutive years with highly different weather in the blooming. Results clearly show that pear does not necessarily produce small amount of nectar as stated in the world literature. In fact, pear can produce extremely high amount of nectar sometimes much higher than other temperate zone fruit trees species but its nectar production is highly subjected to weather, first of all to air temperature. Low nectar production seems to be more frequent than high one and cold weather can prevent its nectar production at all. On the other hand, results corroborate to the earlier statements on the low sugar concentration of pear nectar. There is a highly significant negative correlation between the amount of nectar produced by pear flowers and its sugar concentration (r = -0.52, n = 291, p< 0.001 for 1996, r = -0.34, n = 197, p< 0.001 for 1998). Sugar concentration in individual flowers may be up, to 40% in exceptional cases but generally it is well below 20%. Very high figures for sugar concentration in pear nectar at the literature seem to be incomprehensible. In contrast of some earlier statement in the literature no real difference could be established in the nectar production of pear cultivars, based on much more measurements than in earlier studies. Very low sugar concentration in pear nectar can explain the fact that the overwhelming majority of honeybees are pollen gatherers at pear trees even in the case of exceptionally high nectar production.

Hourly nectar secretion was studied in five pear cultivars between 1997-1999. Some cultivars (e.g. ‘Csákvári téli') secreted nectar continuously during the whole day, offering both nectar and pollen for pollinating insects. Discontinuous secretion (e.g. cvs. ‘Viki', `Nyárig tartó 6/19') is less advantageous from the viewpoint of insect attraction. In some cases, however, discontinuity or continuity of nectar secretion varied even within a cultivar (e.g. 'Solymári cukor', ’Jó szürke’) in different years.

Studies were made on the composition of pollen loads of honeybees captured at the flowers of blooming pear trees in pear plantations. Also the foraging behaviour of honeybees was observed. Overwhelming majority of honeybees visiting the flowers of 13 pear cultivars in 1996 were pollen gatherers (95.6 per cent). Proportion of pure nectar gatherers was as low as some 3.7 per cent and no more than 0.7 per cent performed mixed behaviour. The analysis of pollen loads of bees collected at pear flowers in blooming pear plantations showed that fidelity was as high as 89-90 per cent towards pear, higher than for another fruit species in other studies. Even those plant species that are regarded to be strong competitors of blooming fruit trees in the literature (Taraxacum officinale, Stellaria media, Lamium purpureum) were scarcely represented in the loads. Accordingly, honeybees can be much more important and more effective pollinating agents of pear cultivars than generally believed.

Nectar production of six quince cultivars were measured during three consecutive years. The nectar production of quince can be regarded to be small compared to other temperate zone fruit tree species since quince flowers contained 1.07 ± 0.06 mg of nectar in average. The extreme values, however, ranged between 0.1 and 7.3 mg/flower and this indicated that the nectar production was highly variable. The distribution of the nectar production was definitely skew because low values were definitely much more frequent than the highest ones. Our findings do not corroborate the earlier statements on the high sugar concentration of quince nectar. We found some 21-27% sugar in average, only. The normal distribution of the sugar concentration also indicates that the typical sugar concentration may be between 20-30%. Accordingly, the sugar concentration of quince nectar is rather low compared to other temperate zone fruit tree species (except pear). There was a significant negative correlation between the amount of nectar and its sugar concentration in quince flowers in all of the three years of the study (r= -0.51, n=37, p<0.02 in 1996, r= -0.57, n=28, p<0.1 in 1997, r= -0.35, n=9 I, p<0.001 in 1998). No definite difference was established between the nectar production of quince cultivars. Nevertheless, one cultivar tended to produce less and two other ones produced somewhat more nectar in average than the rest of the 6 cvs investigated but the extreme values of nectar production of cultivars overlapped in most cages.

Studies were made on the bee visitation of 6 quince cultivars and on the foraging behaviour of honeybees at quince flowers for 3 consecutive years. The bee visitation was highly intense because both the plantation and its surrounding was overpopulated by honeybee colonies. Some 5.5 bee visits were counted in average at 50 opening quince flowers in 10 minutes and some 9.7 flowers were visited of the 50 ones observed meanwhile. This equals some 7 bee visits per flower per day per in good weather. Bee visitation, however, was variable and it was greatly different in the three consecutive years with fairly favourable weather. Some cultivars tended to be more and others less visited by honeybees than the others but these differences were not consequent at each occasion. No consequent relationship between the weather and the bee visitation of quince trees could be recognised. It was concluded that .other factors were responsible for the variable nature of the bee activity at quince. Most honeybees tended to collect pollen (51.6% in average for the 3 years) and usually much less gathered for nectar only (19.9%), and the rest of them were mixed behaviour foragers gathering for both (28.5%). There were some slight differences in the foraging behaviour of honeybees at some cultivars but these differences were not always consequent in consecutive years. Also the nectar production of flowers failed to affect the bee visitation and the foraging behaviour of honeybees definitely. For the variable nature of bee visitation and bee behaviour at quince flowers, supplementary pollination is needed to achieve as high set of fruits as high is required to a good crop at quince (at least 20-25% because the flower density is low of this fruit tree species). Since the intensity of bee visitation at the flowers is the only reliable estimate of the necessity of supplementary bee pollination further research is needed to explore the relationship between the number of honeybee visits and the consequent fruit set at quince.

The effect of the partial and/or complete limitation of the insect (bee) pollination period was studied in three consecutive years at 6 quince cultivars. Quince is greatly sensitive to the limitation. Complete limitation of insect pollination resulted in no yield and also a partial limitation of the insect pollination period (exclusion of the bees at the first or at the second half of the blooming) depressed the final set and the yield by 60-70% at least or more, sometimes down to no yield. No real correlation was found between the mean mass of fruits and the fruit set. The mass of fruit seems to be rather a character of the cultivars. A loose but significant correlation was established between the intensity of honeybee visitation and the consequent yield of quince. As much as some 4-5 and 8-10 honeybee visits are needed a day on one flower to achieve the required optimal fruit set that has been declared to be 20-25% for quince in the literature. These are extremely high bee visitation figures, accordingly, no doubt the commercial quince plantations require much higher number of honeybee colonies than other temperate zone fruit tree species to supplementary pollination

The composition of floral nectar in sour cherry cultivars studied in 1997 at Újfehértó was in agreement with our previous data, the three most frequent sugar components being glucose, fructose and sucrose. Nectar secreted at night is generally more diluted than nectar produced during the day. None of the nectar samples produced at night reached the threshold value (100 mg/ml) of bee visitation. In the majority of cultivars the difference in concentration between night and day nectar is not too high, but in two cultivars, 'Korai pipacs' and ‘Újfehértói fürtös’ the difference is quite significant. Most sucrose was found in the nectar of cvs. 'Érdi jubileum' and ‘Újfehértói fürtös’, but a high amount of sucrose was measured also in the flowers of 'Korai pipacs' and 'Meteor USA'. Nectar concentration varies from opening of the flower to petal fall to a smaller or greater degree, depending on the given cultivar. From the 9 sour cherry cultivars studied ‘Újfehértói fürtös’ and Tandy 48' had the most attractive nectar for bees. There was no correlation between nectar composition and free fertilisation. On the basis of nectar composition the majority of the sour cherry cultivars studied can be classified into the sucrose-rich category; only one cultivar, 'Érdi jubileum' had a sucrose-dominant secretory product. The composition of nectar in the studied sour cherry cultivars is preferred by bees.

The percentage of dark staining pollen grains was higher in spring of 1996 than in the previous year. Data in 1998 resemble those of 1995, concerning the large amount of medium staining pollen grains in the majority of clones. Some clones produced excellent quality pollen also in the third year. whereas there were significant differences in other clones in various years.

The warmer February-March period in 1995 induced an early blooming, and frost affected the orchard not only in winter months, but also immediately before and during blooming. Thus, frost was the possible cause of weaker quality pollen this year. In 1996 warming began a bit late, but it was not broken by drastic falls in temperature, except for the middle of April, when a smaller frost affected the orchard. It is likely that this frost did not influence pollen quality of `Besztercei' and 'Early Besztercei' plum clones significantly. In 1998 warming was continuous and steady, the orchard was not affected by frost immediately before blooming. In March, however, there was frost almost every day, according to daily minimum temperatures.

Observations were made at two growing sites, Siófok and Szatymaz, in the years 1998 and 1999, on 16 peach varieties. The production of nectar was measured, the foraging behaviour of bees, fruit set and the effect of exclusion of bee visits for different periods were observed systematically.

Production of nectar confirmed earlier data, 9.09 mg per flower in average. There was large variation due to variety and date of observation. Bee visits were relatively abundant. At favourable weather, 1 to 30 visits/flower/day occurred in the average. Artificial hand pollination increased fruit set, substantially. Open pollination yielded superior fruit set than self pollination, without bees. Supplementary bee pollination can be regarded to be beneficial to peach production as well.

Five pepper cultivars were mechanically inoculated with isolates of three Tobamovirus species, viz. the "Gelb" strain of tobacco mosaic virus (TMV-G), the XM-isolate of dulcamara yellow fleck virus (DYFV-XM) and the Nov/H isolate of pepper mild mottle virus (PMMV-Nov/H), respectively. Symptoms caused by the viruses were characterised. The viruses were sucessfully re-isolated from different organs (roots, leaves, fruit parts) of susceptible peppers to test plants. It was estabilished, that the pollen of diseased peppers carried infective virions at least on their surface. Washes of seeds were highly infective, but no infectivity was found after treatment of the seeds with 2% NaOH or 10% Na3PO4. No infectivity of inocula prepared from seed-coats of alkaline treated seeds was established. Infection of young seedlings grown from untreated seeds was demonstrated, while the seedlings came from alkaline treated seeds remained free of infective virus. The possible role of pollen and seed in the epidemiology of Tobamoviruses pathogenic to pepper as well as the importance of seed treatment is discussed.

The Apiaceae family consists of several species which are well known for their therapeutical use and also as spice plants. Although fennel (Foeniculum vulgare Mill.), caraway (Carton carvi L.), anise (Pimpinella anisum L.), coriander (Coriandrum sativum L.), dill (Anethum graveolens L.), angelica (Angelica archangelica L.) and lovage (Levisticum officinale Koch.) are also economically considerable cultures, data on their flowering biology are rather scarce. This review supports data on the characteristic constitution of flowers and inflorescences, flowering dynamics, pollination mechanism and crossability of some of the most significant medicinal and spice species.

The inflorescence is a compound umbel. Flowers are hermaphrodite, however also monoeceous, mainly male flowers are turning up too. In blooming of a plant individual, a strict sequence is observed. The main umbel is the first to bloom, followed by the different range umbels in order of their range. In their pollination, both wind and insects are considered to be active. For each species, proterandry is characteristic. Although autogamy is considered to be almost unpossible, and geintonogamy as well as xenogamy to be the most characteristic ways of fertilization, several cases of self-fertilization proved to be also successful. Crossability among cultivars or species is depending on the genotype combination and usually produces less seed set than the above forms of fertiliiation.

Several species in the Lamiaceae family are therapeutically significant medicinal plants. Such as sage (Salvia officinalis L.), rosemary (Rosmarinus officinalis L.), peppermint (Moldier piperita L.). marjoram (Majomna hortensis Winch), thyme (Thymus vulgaris L.), Origanum spp., hyssop (Hyssopus officinalis L.), Lavandula spp., basil (Ocimum basalicum L.), lemon-balm (Melissa officinalis L.) are regularly produced in Hungary. Nevertheless, data on their floral biology are not satisfactory. This review intends to gather information on the characteristic constitution of flowers and inflorescences, blooming dynamics, pollination mechanism and crossability of some of the most significant medicinal and spice species

Flowers are mainly hermaphrodite, however, also monoeceous ones are turning up: pistillate flowers marjoram, tyme, oregano. In flowering of a plant individual, a basipetal blooming sequence is described. These species have a floral structure fully adapted to pollination by insects, and the majority of them is an excellent honey-plant (lavandel, hyssop, sage, etc.).

Flowers exhibit characteristic proterandry, therefore autogamy is considered to be almost impossible, and geintonogamy as well as xenogamy to be the most characteristic ways of fertilization. However, in several cases self-fertilization proved to be also successful. Crossability among cultivars or species is depending on the genotype combination and usually produces less seed than the above forms of fertilization.

In the present experiment pollen viability of snowdrop were tested using four vital dyes (TTC, Baker' procedure, MTT and FDA) to determine their potential to differentiate fresh living pollen from pollen heated for 12 hours at 80 °C (killed pollen). We found that two of the four dyes previously employed to determine viability also stained killed pollen in the case of snowdrop, while the FDA and MIT did not. We suggest that the two latter are probably the best method to test snowdrop pollen viability, since they do not normally stain either killed or aborted pollen.

Differences in the attractiveness of pollen according to plant families and species for the honeybees had been examined by researchers, but pollen samples of cultivars were not tested yet. This examination attempts to evalu2te the differences in attractiveness of pollen of some cultivated and ornamental apples to honeybees. We applied biotest with equal amounts of hand collected pollen samples and by a nucleus of bees with brood. The evaluation was based on the number of visitation by foraging honeybees on the samples during the time of the experiment which was determined. Pollenkitt samples were also tested in a separate experiment.

The data were analysed by variance analysis and significant differences were found according to the number of bee visitation in the attractiveness of the pollen and pollenkitt samples of apple cultivars.

The visualisation of pollenkitt was done by light microscope and SEM.

The model of a pollen mixing device was tested in sunflower culture in order to detect the effectiveness in helping the out-going honey bees to carry more and different varieties of pollen grains on the body and body hairs. This model is build from line hairs of brushes facing each other that can keep and release pollen grains. The brushes are functioning as pollen receiver-transmitter units. The effectiveness was tested by examining the number and the species of the pollen grains adhered to the bees passing through the entrance. The amount of pollen and the number of pollen species on the hive with the device was significantly higher than on the control hive.

The data were analysed by variance analysis and significant differences were found according to the number and species of pollen grains between the incoming and outgoing bees.

Honeybee races and ecotypes of different genetic background have different population development in spring. Some of them can reach the necessary population size by the beginning of Robinia pseudoacacia (black locust) blooming period. There were significant differences in the spring population development between the colonies of different genetic background. The Italian races (A. m. ligustica) and their cross-breeds over-wintered poorly in Hungary, their spring population was low and they collected small amount of Robinia honey. The Austrian improved Carniolan (A. m. carnica) colonies over-wintered well, they had the largest spring population in both years. There was no significant difference between the size of the spring population of the same colonies of different genetic background in 1995 and 1996. The rate of the population development of the colonies was different in the two examined years. There was strong correlation (r = 0.8) between the spring population size and the Robinia honey yield, and between the mid-April population size and the Robinia honey yield of the colony groups of different genetic background. Spring population size also important in the effective pollination of fruit tree species that bloom earlier than the black locust trees.

In the past decades fruit production has become more and more intensive and so the interest towards the bee pollination of fruit trees has increased in Hungary. Thus insect pollination research has become more intensive in the eighties and in the nineties. Recently, high density fruit orchards are planted of dwarf trees at more and more farms in Hungary and these need highly controlled growing practices involving insect pollination, too. This is a new challenge of insect pollination research in Hungary. So it seems to be worth to compile a list on research publications having been appeared so far to create a basis to the ongoing new research. Besides papers strictly on the insect pollination of fruit trees publications on some closely related topics, first of all on the nectar production and on other flower characteristics of fruit tree cultivars that directly influence bee activity have also been included on the list.