The effect of nectar production to the gathering behaviour of honeybees and to the foraging activity of wild bees at apple flowers

Intensity of bee visitation (honeybees and wild bees), foraging behaviour of honeybees and nectar parameters (nectar production, sugar concentration, sugar content in nectars) were inspected at 18 apple cultivars for three consecutive years. Honeybee visitation was calculated to be some 3.07 bee visits at a single apple flower a day but wild bee activity was very low, only some 0.11 wild bee visits per flower per day.

The nectar production (nectar content) of apple flowers was fairly different according to the cultivars and the nectar production (nectar content) of flowers was negatively correlated with the sugar concentration in nectar. Interestingly, at the majority of the inspected 18 cultivars the nectar production has shown minor or no changes in the morning and at the afternoon. The nectar production (nectar content) of flowers clearly encouraged the total bee activity at the inspected cultivars (r = 0.54). Bees visited abundant nectars with greater sugar concentration most intensely than lees abundant nectars with smaller sugar concentration.

Nectar parameters, however, affected the activity of honeybees with different types of gathering behaviour in different way. More intense activity by pollen gatherer and mixed behaviour honeybees was observed at cultivars producing abundant nectar (pollen gatherers: r = 0.65, mixed behaviour: r = 0.79). The activity of pollen gatherer honeybees and of mixed behaviour bees, on the other hand, was negatively correlated to the sugar concentration in nectar (pollen gatherers: r = -0.51, mixed behaviour: r = -0.73). This can be explained by the fact that their behaviour was much more affected by the amount of pollen than by any nectar parameters. Accordingly, "mixed behaviour honeybees" should probably rather be called as "pollen gatherers with nectar load" instead of "nectar gatherers with pollen load", as widely used in literature. The activity of ineffective side worker nectar gatherers was greatly encouraged by the sugar concentration in nectar (p = 0.63), similarly to the pure (top) nectar gatherer honeybees (r = 0.72). There was a negative relationship between the nectar production (nectar content) of flowers and the activity of these behaviour classes (nectar gatherers: r = -0.47, side workers: r = -0.36).

Concluding the findings we can state that the activity of pollen gatherers and mixed behaviour honeybees is strongly encouraged by greater nectar production (greater nectar content) of apple flowers. Their activity, however, is mostly dependent firstly on the amount of pollen. This is the reason why the sugar concentration is in a negative relationship to the activity of pure pollen gatherers and mixed behaviour bee. On the other hand, pure nectar gatherers and side worker nectar gatherers are greatly encouraged by the sugar concentration of apple nectar the amount of nectar was not a definite influence on their activity.

Side worker honeybees occurred at each cultivar inspected, however, their ratio varied widely among cultivars. So we can draw the conclusion that, in contradiction with earlier believes, side worker nectar gathering is a general phenomenon at apple flowers. The activity of side workers depends first of all on the relative position of stamens and petals; however, low sugar concentration of nectars can probably moderate their activity but probably do not affect pollen gatherers and mixed behaviour bees.

Wild bee visitation was very low; the wild bee species observed (Osmia cornuta, Andrena flavipes, Anthophora acervorum, Bombus terrestris, Bombus lapidarius and Halictus simplex) comprised some 3.44 per cent of the total bee visitation at apple flowers only. Wild bee visitation was in a positive correlation with the nectar production (nectar amount) of flowers (r = 0,4) and with the sugar content of nectar (r = 0.46) but it was negatively correlated with the sugar concentration in nectar (r = -0,27).

Our result calls the attention to the importance of nectar parameters in the effective honeybee pollination of apple plantations. Most effective pollinating activity of honeybees can be expected in plantations with cultivars of high nectar production because this feature seem so encourage the activity of those behaviour classes (pure pollen gatherers and mixed behaviour bees) that are the most effective pollinating agents among honeybees.

Flower constancy of honeybees (Apis mellifera L.) to blooming pear plantations

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.


Flower characters and self-fertilization capacity in relation to the bee pollination at sour cherry cultivars

Detailed studies and comparisons were carried out on those flower characters of sour cherry cultivars that may affect bee pollination of flowers. Flower characters of sour cherry are fairly similar to other temperate zone fruit tree species. Their relatively small flowers distinguish the Cigánymeggy-types of cultivars from the flowers of tart cherries cultivars that are conspicuously larger, almost as large as the sweet cherry flowers. The relative position of flower organs was much more variable according to the season than according to the cultivars. So the differences were rather the consequences of seasonal effects than of variety features of sour cherry cultivars. As far as individual cultivars are concerned differences in the nectar production and the sugar concentration are revealed rather between groups of cultivars than between individual cultivars. The pollen production of flowers was extremely changeable in consecutive years. Most honeybees collected nectar at sour cherry flowers; pure pollen gatherers and mixed behaviour bees were half as frequent but differences among the behaviour of honeybees according to cultivars cannot be stated. The fidelity of honeybees to sour cherry is less expressed than to some other fruit tree species. Accordingly, it is very strongly suggested to take the competitive effect other plant species (weeds) flowering in and around the orchard carefully into account when organizing additional bee pollination in sour cherry plantations. Several sour cherry cultivars possess more or less self-fertilization capacity but this is greatly changeable according to the season. It has been proved that self-sterile sour cherry cultivars are sensitive even on the partial restriction of the effective time of bee pollination and it is to be stressed too that even in the case of partly self-fruitful cultivars bee pollination is also vital in yield formation because medium or strong restriction of the effective bee pollination period is of a definite negative effect on their fruit set and yield. In years with unfavourable weather the yield can dramatically be reduced sometimes down to nil. However, very high fruit set is also unfavourable because a negative correlation was detected between the final set and the mean mass of fruits.

Effect of intensity of bee visitation and the foraging behaviour of honeybees on the fruit set and yield of apple (Malus domestica Borkh.)

Based on the results of our experiments, both the relative and the effective intensity of bee visitation were rather different depending on the cultivars as well as the time of the day. While it varied between relatively extreme values (40-80%) in the morning for the varieties examined, there were much smaller differences between the intensity of bee visitation at the afternoon, because the relative bee visitation attained 70-90% at each cultivar. These results showed that the differences arising from intensity of bee visitation of different cultivars should be taken into consideration more carefully in the morning in orchard planning and in estimating the number of honeybee colonies required.

The results showed that the greatest percentage of fruit set and the highest number of viable seeds per fruit were measured on branches of those cultivars that were most frequently visited by pure pollen gatherer bees as well as by bees collecting both nectar and pollen (mixed behaviour). The effect of pure pollen gatherers and of bees with mixed behaviour was highly significant from the statistical point of view on the fruit set and the number of viable seeds per fruit.

Those bees that were sucking nectar only from apple flowers did not proved to be effective pollinators at all. Relationship between their number and the fruit set as well as the number of viable seed per fruit were not significant because the coefficient of correlation was close to nil.

The ratio of side worker nectar gatherers was negatively correlated with the fruit set and the seed content of fruits of apple cultivars examined at both of our experimental sites, at Mosonmagyaróvár and Feketeerdő as well. The presence of side worker nectar gatherers resulted in higher decrease of fruit set and seed content of fruits at Feketeerd6 than at Mosonmagyaróvár, especially in the morning.

The effect of flower visiting intensity by other pollinating insects was found to be fairly variable according to the time of the day. In the morning they had no effect on the fruit set as well as on the seed content of fruits either at Mosonmagyaróvár or at Feketeerdo. However, at the afternoon, when their intensity was greater, the correlation was a bit stronger.

Nectar production of pear (Pyrus communis L.) cultivars

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.


Bee pollination and association of apricot varieties

Apricot yields are highly variable according to the season. The variation is caused mainly by the adversities during the critical processes of floral biology, i.e. blooming and fertilisation. On the basis of information concerning blooming time and mutual compatibility relations of apricot varieties a system of securing regular and adequate yields has been developed.

Winter frosts of the continental type are well tolerated by most of the apricots, however, after the end of rest period, flower buds are loosing frost tolerance, 'rapidly.

Being one of the fruit species blooming earliest during the early spring, apricot start to bloom in Hungary around the end of March or early April as a mean of many years, but it also happened, exceptionally that apricot started to bloom at February 20 (at Letenye South Hungary). The early season, exposes the floral organs to frost injuries. As a consequence, apricot orchards on the Great Plain produce low yields in 3 years, intermediate yields in other 3 years out of a ten-year-period.

Moreover, weather conditions during the blooming period are often unfavourable for pollination. Cool, windy and rainy weather prevents the flight of insects and on the other hand, warm spells shorten the blooming process, nectarines and stigmata get dry and the female gametes loose viability before effective pollination occurres.

The fertility of individual cultivars are meeting different obstacles. Apricot cultivars differ greatly in the rate of flowers bearing underdeveloped pistils, which may attain even 60% (e.g. Orangered). New commercial cultivars are often self-incompatible. Local varieties of that type in Hungary are the „óriás" varieties (e.g. Ceglédi óriás, Szegedi mammut), and the new hybrid Ceglédi Piroska. Many of the cultivars are variable in their self-fertility (partially self-fertile): Budapest, Harmat, Korai piros, Mandulakajszi.

Inter-incompatibility is also known in apricots. The „óriás " varieties do not fertilise each other. During the growth of fruits, cool spells (2-4 °C) caused severe fruit shed in Ceglédi óriás.

Apricot flowers produce pollen and nectar at average rates related to other fruit species, thus bees are attracted sufficiently. Bee visits are very variable according to growing site and season. Most of the bees are pollen gatherers but sometimes nectar suckers are in majority. Bee pollination is necessary not only for the self-incompatible varieties but also to enhance the yield of self-fertile varieties.

Taking the blooming and fertility relations of the cultivars into account, plantations should not exceed two rows to a particular self-incompatible varieties, and possibly two different polliniser varieties are suggested to be planted as flanking the block in question.

In commercial plantations 2 to 4 bee colonies per hectare are proposed to move for the whole blooming period.