Why be a honeyless honey mesquite? Reproduction and mating system of nectarful and nectarless individuals

Populations of Prosopis glandulosa var. torreyana in the Chihuahuan desert have a fixed dimorphic system of nectar production in which half the individuals produce nectar (are nectarful) and the other half are nectarless. We analyzed the impact of nectar production on different estimates of fitness, comparing nectarful against nectarless individuals in size, mating system, seed traits, and fruit set in a 1-ha scrubland. Of the reproductive individuals (358), 46% were nectarful and 54% were nectarless. Neither tree size nor flowering phenology differed between nectar morphs. Fixation indices (F) for both progeny (F = −0.2) and adults (F = −0.45) were negative, and high heterozygosities were found in adults and progeny (H = 0.45). No differences were found between nectar morphs for F, H, and single (t_s = 1.1) and multilocus (t_m = 1.03) outcrossing rates. Controlled pollinations showed differences between selfing and control treatments with no differences between nectar morphs. Nectarless individuals produced significantly more pollen grains than did nectar producers, but all other measured floral traits showed no differences. Nectarful trees were visited by pollinators 21 times more often and had a significantly higher overall fruit set than did nectarless trees. No differences between nectar morphs in seed mass or in percentage seed germination were found, but heavier seeds tended to have higher heterozygosities. Both morphs had similar success as females, but nectarless trees had ∼7% higher male function. We discuss three possible scenarios for the evolution of the fixed dimorphism in nectar production, two involving unstable phases (substitution of one morph by the other, and evolution towards dioecy) and one stable scenario (maintenance of the dimorphic system).     

Bee diversity on nectarful and nectarless honey mesquites

Nectar production has been proposed as an adaptation to attract pollinators that benefit from this resource. Energetic investments may be expensive, so some species such as Prosopis glandulosa have developed a dimorphic system of nectar production, which is expected to affect floral visitor behaviour and then plant fitness. We quantified bee diversity during a 2 year period in a population of the honey mesquite in order to determine changes in bee diversity due to the presence of nectar, bee preferences to collect either nectar of pollen, and to determine between year variations of bee faunas. Floral visitors were captured at three different times of the day during the flowering seasons of 1994 and 1995, in a population of Prosopis glandulosa which has a 1:1 proportion of nectar: nectarless individuals. Pollinators were clearly distinct between nectar morphs, bee species diversity and relative abundance of visits were significantly greater on nectarful than on nectarless plants, with species on nectarless individuals being a subset of those in the nectarful morph. Our results suggest differences in the function of floral rewards (i.e., nectar and pollen) to attract floral visitors. For the Chihuahuan arid environment, mesquite provides floral rewards with ease, quantity and quality for close to 10% of all bee fauna making them important components of these communities.     

Stem demography of Prosopis glandulosa var. torreyana in vegetation arcs and associated bare areas

Abstract. Vegetation arcs are characterized by dense patches of up to 100% plant cover surrounded by bare areas where cover is around 5%. These patterns of contracted vegetation have been reported mainly in alluvial fans of arid and semiarid zones (known as bajadas) with gentle slopes (0.2–2%), summer rainfall, and heavy showers that run as sheet-flow. It has been suggested that run-off water from bare areas is stopped at the front of the arcs and, depending on the amount of rainfall, may advance superficially into the arc, reaching the downslope limit less often. These dynamics would imply a gradient of water availability inside the arc, higher at the upper edge. We termed this the moisture gradient hypothesis. We indirectly tested this hypothesis by following the demography of a cohort of stems of the honey mesquite Prosopis glandulosa var. torreyana in three vegetation arcs and their corresponding bare areas during a yearly cycle at the Bolsón de Mapimí in the Chihuahuan Desert. We selectively removed different plant life-forms (trees, shrubs and herbs) in three positions within the arcs; no manipulation was possible in the bare areas due to low plant cover. The results suggested that removal of other plant life-forms, mainly grasses, affected growth and survival of mesquite low (but not high) stems, making them accessible to medium-sized mammal browsers (jackrabbits and packrats). These results suggest that, at least for the honey mesquite, water availability in vegetation arcs was not the most strongly limiting factor for shoot performance and demography, and that browsing damage, a biotic constraint, took precedence over resource limitation.     

BIOMASS AND NET PRIMARY PRODUCTION OF PROSOPIS GLANDULOSA (FABACEAE) IN THE SONORAN DESERT OF CALIFORNIA

Prosopis glandulosa var. torreyana accounts for nearly 90% of the total plant cover in a mesquite woodland community near Harper's Well along the southern margin of the Salton Sea in the Sonoran Desert of California. Total above-ground biomass in ten individuals studied in detail ranged from 43–760 kg per plant and 1.9–8.5 kg m^-2 canopy area. Stand biomass ranged locally from a high of 23,000 kg ha^-1 near the wash to 3,500 kg ha^-1 in the fringe of this mesquite stand. Net above-ground primary production for 1980 had a mean of 2.2 kg m^-2 canopy for shrub forms and 5.3 kg m^-2 canopy for tree forms. Mean Prosopis stand production for 1980 was 3,650 kg ha^-1, an extremely high value for desert communities. This level of production is particularly high in relation to the low mean annual precipitation of approximately 70 mm. New woody tissues in trunk and branches accounted for 51.5% of the allocation of productivity in Prosopis, a remarkably high woody allocation for a desert plant. Only 33.6% of net primary production was allocated to leaves.     

Cerambycid girdling and water stress modify mesquite architecture and reproduction

This study evaluated the effect of an outbreak of the cerambycid beetle, Oncideres rhodosticta, on branch growth and inflorescence production of the mesquite Prosopis glandulosa var. torreyana, and on larvae mortality in girdled branches at two sites (dry and wet) in the southern Chihuahuan Desert of Mexico. We compared stem growth responses to girdling in branches of similar sized trees from both sites over 42 months. The number of larvae per girdled branch was similar between sites, indicating similar ovipositing effort regardless of water stress. However, the proportion of dead larvae was significantly lower in trees at the dry site. On average, girdling reduced 96% of the stem length at both sites. At the end of the first year, 25% of the original stubs survived at the dry site, compared to 90% at the wet site. Girdling also broke apical dominance and allowed for the development of lateral buds from the surviving stubs, which produced many fewer branches and inflorescences at the dry site compared to the wet site, where a compact crown was developed. Water stress and girdling have a combined effect on mesquite architecture and reproduction, since more stubs died at the dry site while new branches from surviving stubs developed at the wet site, recovering the original lost biomass but changing the appearance of the tree.     

Accumulation of human EGF in nectar of transformed plants of Nicotiana langsdorffii x N. sanderae and transfer to honey by bees

Honey has been used successfully in wound healing for thousands of years. The peptide hormone human epidermal growth factor (hEGF) is also known to have a beneficial effect in various wound healing processes via mechanisms that differ from those for honey. In this study, we show that hEGF can be incorporated into honey via nectar. Plants of Nicotiana langsdorffii × N. sanderae were transformed with the gene for hEGF, equipped with a nectary‐targeted promoter and a signal sequence for secretion to nectar. These plants accumulated hEGF in the nectar. The maximum hEGF concentration recorded with ELISA in these plants is 2.5 ng·ml^?1. There is a significant linear relationship (P < 0.001) between hEGF concentration and induction of hEGF‐receptor phosphorylation. Since the flower morphology of these plants did not allow production of honey from their nectar, we used feeding solutions, spiked with synthetic hEGF, to study transfer of this peptide into honey through bee activity. Transfer of hEGF from a feeding solution to honey by bees occurred with retention of the hEGF concentration and the capacity to induce hEGF‐receptor phosphorylation. These observations indicate that plants can function as a production platform for honey containing biologically active peptides, which may enhance wound healing and other biological processes.     

Use of thermocouple psychrometry in field studies of soil-plant-water relationships

Summary Use of Spanner thermocouple psychrometers to measure soil water potentials and internal water potentials of woody plants was studied using honey mesquite (Prosopis glandulosa var. glandulosa) growing under two soil water regimes. A high correlation (r=0.92 or greater) existed between the soil water potential (bars) measured with Spanner thermocouple psychrometers and soil water content (percent) obtained from gravimetric samples on a fine sandy loam soil. The influence of soil water potential and diurnal fluctuations were easily measured in honey mesquite growing under two soil water regimes. The results of this study indicated that thermocouple psychrometers can effectively be used in field studies involving soil-plant-water relationships. College of Agricultural Sciences Publication Number T-9-130. College of Agricultural Sciences Publication Number T-9-130.     

Evidence for mutualism between a flower-piercing carpenter bee and ocotillo: use of pollen and nectar by nesting bees

Abstract.

• 1 Carpenter bees (Xylocopa californica arizonensis) in west Texas, U.S.A., gather pollen and ‘rob’ nectar from flowers of ocotillo (Fouquieria splendens). When common, carpenter bees are an effective pollen vector for ocotillo. We examined ocotillo's importance as a food source for carpenter bees.
• 2 The visitation rate of carpenter bees to ocotillo flowers in 1988 averaged 0.51 visits/flower/h and was 4 times greater than that of queen bumble bees (Bombus pennsylvanicus sonorus), the next most common visitor. Nectar was harvested thoroughly and pollen was removed from the majority of flowers. Hummingbird visits were rare.
• 3 Pollen grains from larval food provisions were identified from sixteen carpenter bee nests. On average, 53% of pollen grains sampled were ocotillo, 39% were mesquite (Prosopis glandulosa), and 8% were Zygophyllaceae (Larrea tridentata or Guaiacum angustifolium). Carpenter bee brood size averaged 5.8 per nest.
• 4 We measured the number of flowers, and production of pollen and nectar per flower by mature ocotillo plants, as well as the quantity of pollen and sugar in larval provisions. An average plant produced enough pollen and nectar sugar to support the growth of eight to thirteen bee larvae. Ocotillo thus has the potential to contribute significantly to population growth of one of its key pollinators.
• 5 Although this carpenter bee species, like others, is a nectar parasite of many plant species, it appears to be engaged in a strong mutualism with a plant that serves as both a pollen and as a nectar source during carpenter bee breeding periods.

     

Influence of soil temperature on niacin and thiamine in honey mesquite

Summary The influence of soil temperature was examined on niacin and thiamine concentration in honey mesquite (Prosopis glandulosa var.glandulosa) seedlings. The seedlings were grown in soil temperature regimes of 21, 27, and 32°C in a controlled environment growth room. Nodulation randomly occurred on the roots of the seedlings, necessitating separate analysis according to the occurrence of nodulation. Roots of nodulated seedlings from the 21°C soil temperature regime contained greater quantities of niacin and thiamine compared to root samples from seedlings grown in either 27 or 32°C regimes. Niacin concentration of non-nodulated seedlings was highest in samples from seedlings grown in the 27°C soil temperature regime and lowest in samples from seedlings grown in the 21°C regime. Thiamine concentration was the greatest from non-nodulated seedlings grown in the 27°C soil temperature regime, while the thiamine concentration of non-nodulated samples from the 32°C regime was the least. Optimal soil temperature for honey mesquite root growth appears to be about 27°C. At sub-optimal soil temperatures niacin might have limited ‘growth’ while at supra-optimal soil temperatures, thiamine might be a limiting factor. College of Agricultural Sciences Contribution No. T-9-164.     

Bees get a head start on honey production

Nectar concentration is assumed to remain constant during transport by honeybees between flowers and hive. We sampled crop contents of nectar foragers on Aloe greatheadii var. davyana, a major winter bee plant in South Africa. The nectar is dilute (approx. 20% w/w), but the crop contents of bees captured on flowers are significantly more concentrated. In returning foragers, the concentration increases further to 38–40%, accompanied by a volume decrease. The doubling of sugar concentration suggests that nectar is regurgitated onto the tongue and evaporated during foraging and on the return flight. Processing of the dilute nectar into honey thus begins early, aided by low ambient humidities. This has implications for honeybee thermoregulation, water balance and energetics during foraging, and for the communication of nectar quality to recruits.     

Reproductive biology of Alpinia blepharocalyx (Zingiberaceae): another example of flexistyly

The floral biology and breeding system of Alpinia blepharocalyx were studied in Yunnan province, southwest China. Our results indicate that populations of A. blepharocalyx have two floral morphs, which differ in flowering behaviour: the cataflexistylous morph in which the stigma is held erect above the dehiscent anther when anthesis begins in the morning and becomes curved under the anther at afternoon, and the anaflexistylous morph in which the receptive stigma is curved under the indehiscent anther first and moves into a reflexed superior position above the anther as it begins to shed pollen in the afternoon; the flowering period of both floral morphs is 12 hours; the stylar movements in the two floral morphs are synchronous, and they have similar traits to those found in other Alpinia species previously reported. Also, on average, cataflexistylous flowers are larger than anaflexistylous flowers, especially the labellum and corolla tube length; moreover, the P/O ratio of the two floral morphs is significantly different: the cataflexistylous morph has more pollen grains and fewer ovules than the anaflexistylous morph; the evolutionary significance of this comparison is discussed. Cataflexistylous flowers were observed to produce a lower concentration of nectar than that of anaflexistylous flowers before 11:00 a.m., but they then have higher nectar concentration; cataflexistylous flowers always have more nectar than anaflexistylous flowers. Flowers of A. blepharocalyx were visited by fewer insects at a lower frequency, including honey bees (Apis cerana cerana) and two species of carpenter bees (Xylocopa spp.). No significant differences were found in the fruit set among the hand-pollination treatments regardless of selfing (geitonogamy and autogamy) or crossing (intermorph and intramorph); but all hand-pollination treatments exhibited much higher fruit set than the controls; meanwhile, no fruit set occurred in the unpollinated bagged plants. Pollen tube growth was examined using fluorescence microscopy following self, intramorph, and intermorph hand pollinations at different times of the day; the pollen tubes of both floral morphs have the same growth rate and the capability to reach the ovary. Both the results of pollen tube growth and fruit sets under different pollination conditions show that A. blepharocalyx is self-compatible and dependent upon insects for pollination. The distinctive flexistyly mechanism of A. blepharocalyx is likely a floral mechanism that promotes effective intermorph pollen transfer.     

Postpollination Nectar Reabsorption and Its Implications for Fruit Quality in an Epiphytic Orchid1

We present evidence that pollination triggers nectar reabsorption in flowers of the epiphytic orchid Mystacidium venosum. The amount of sugar in nectar of M. venosum decreased significantly by more than 50 percent within 72 hours of pollination. Hand–pollinated flowers from which nectar was previously removed set significantly smaller fruits with a lower percentage of viable seeds than hand–pollinated flowers containing nectar, suggesting that resources reclaimed by nectar resorption are allocated to fruit production.     

Regulation of nectar collection in relation to honey storage levels by honey bees, Apis mellifera

Honey bees collect distinct nutrient sources in the form ofnectar (energy) and pollen (nitrogen). We investigated the effectof varying energy stores on nectar and pollen foraging. We foundno significant changes in nectar foraging in response to changesin honey storage levels within colonies. Individual foragersdid not vary activity rates or nectar load sizes in responseto changes in honey stores, and colonies did not increase nectarintake rates when honey stores within the hive were decreased.This result contrasts with pollen foraging behavior, which isextremely sensitive to colony state. Our data show that individualforaging decisions during nectar collection and colony regulationof nectar intake are distincdy different from pollen foraging.The behavior of honey bees illustrates that foraging strategy(and therefore foraging models) can incorporate multiple currencies,including both energy and protein intake.[Behav Ecol 7: 286–291(1996)]     

Nutritive composition of green and ripe pods of honey mesquite (Prosopis glandulosa,fabaceae)

Pods from Prosopis glandulosa (honey mesquite) were harvested at seven different growth stages. Protein content (N × 6.25) of the whole pod decreased with maturation with small differences occurring after stage 3. Seed contained about 82% of the total pod protein. Carbohydrate accumulated primarily in the pericarp, and ripe pods (stage 7) contained about 80% total carbohydrate. Fat content of the pod increased slightly with maturation while total ash and fiber (ADF) decreased. Small differences existed between stages 4 and 7 in the essential amino acid pattern of whole pods. The total sulfur-containing amino-acids were the most limiting in both growth stages; the pericarp protein had a relatively better amino-acid pattern than did the seed protein. Both green and ripe honey mesquite pods provide similar amounts of micro- and macro-minerals.     

Introducing plants for bee-keeping at any cost? – Assessment of Phacelia tanacetifolia as nectar source plant under xeric Mediterranean conditions

 We assessed nectar and honey potential of Phacelia tanacetifolia, an American plant, under Mediterranean conditions. Its flowering occurred during the major flowering season unless intensely irrigated, whereas duration and flower life span were shorter compared to continental conditions. Mediterranean climate limitations to nectar secretion were assessed on plants grown under natural conditions (xeric) vs. regular irrigation. May flowering xeric plants produced less nectar per flower than the irrigated ones, but had the same nectar potential per surface area. On the contrary, at the cost of intense irrigation, July flowering plants produced much higher nectar per flower and per surface area compared to xeric ones. In all flowering periods and sections honeybees were the most constant and numerous visitors, visiting the flowers mainly for nectar, whereas solitary bees were scarce. Based on our results, we suggest that although the plant may be a good nectar source for honeybees in some cases, we have serious reservations for a generalised use in the Mediterranean. Received September 3, 2002; accepted November 18, 2002 Published online: June 2, 2003     

Floral nectar production: what cost to a plant?

Floral nectar production is central to plant pollination, and hence to human wellbeing. As floral nectar is essentially a solution in water of various sugars, it is likely a valuable plant resource, especially in terms of energy, with plants experiencing costs/trade-offs associated with its production or absorption and adopting mechanisms to regulate nectar in flowers. Possible costs of nectar production may also influence the evolution of nectar volume, concentration and composition, of pollination syndromes involving floral nectar, and the production of some crops. There has been frequent agreement that costs of floral nectar production are significant, but relevant evidence is scant and difficult to interpret. Convincing direct evidence comes from experimental studies that relate either enhanced nectar sugar production (through repeated nectar removal) to reduced ability to produce seeds, or increased sugar availability (through absorption of additional artificial nectar) to increased seed production. Proportions of available photosynthate allocated by plants to nectar production may also indicate nectar cost. However, such studies are rare, some do not include treatments of all (or almost all) flowers per plant, and all lack quantitative cost–benefit comparisons for nectar production. Additional circumstantial evidence of nectar cost is difficult to interpret and largely equivocal. Future research should repeat direct experimental approaches that relate reduced or enhanced nectar sugar availability for a plant with consequent ability to produce seeds. To avoid confounding effects of inter-flower resource transfer, each plant should experience a single treatment, with treatment of all or almost all flowers per plant. Resource allocation by plants, pathways used for resource transfer, and the locations of resource sources and sinks should also be investigated. Future research should also consider extension of nectar cost into other areas of biology. For example, evolutionary models of nectar production are rare but should be possible if plant fitness gains and costs associated with nectar production are expressed in the same currency, such as energy. It should then be possible to understand observed nectar production for different plant species and pollination syndromes involving floral nectar. In addition, potential economic benefits should be possible to assess if relationships between nectar production and crop value are evaluated.     

Flower visitation patterns of the coexisting honey bees Apis cerana japonica and Apis mellifera (Hymenoptera: Apidae)

We compared flower visitation patterns of two coexisting honey bees, Apis mellifera Linnaeus and Apis cerana japonica Radoszkowski, on 20 plant species, including three exotics, under natural conditions in Nara, Japan, from April to August 2012. We also measured flower color based on bee color vision (15 flower species), nectar volume (nine species) and nectar concentration (eight species). Flowers colored white, pink, red, purple and cream were classified as bee‐blue‐green, and yellow was classified as bee‐green. Apis cerana visited 14 plant species and A. mellifera visited 11. Although the two Apis species are similar in morphology, they visited different plants: in particular, A. cerana visited native plant species more often than did A. mellifera. Both A. mellifera and A. cerana visited not only nectariferous flowers but also those with no nectar. We also found different visitation patterns between A. cerana and A. mellifera: Apis cerana more often visited flowers with smaller color angle (bee‐blue‐green), lower chroma and higher brightness, and flowers secreting nectars of higher concentration and smaller volume than did A. mellifera.     

Influence of adjuvants and plant growth regulators on herbicide performance in honey mesquite

Addition of surfactant I (trimethylnonylpolyethoxyethanol) or surfactant II (4-isopropenyl-1-methyl-cyclohexane) at 1.0 and 0.6% (v/v) of the spray solution enhanced the phytotoxicity of clopyralid (3,6-dichloro-2-pyridinecarboxylic acid), the triethylamine salt of triclopyr {[(3,5,6-trichloro-2-pyridinyl)oxy]acetic acid}, picloram (4-amino-3,5,6-trichloro-2-pyridinecarboxylic acid), and 2,4,5-T [(2,4,5-trichlorophenoxy)acetic acid)] applied at 0.14 kg ae/ha to greenhouse-grown honey mesquite (Prosopis glandulosa Torr.). Application of benazolin [4-chloro-2-oxo-3(2H)-benzothiazolacetic acid] increased the phytotoxicity of all herbicides, but ethephon [(2-chloroethyl)phosphonic acid] and mefluidide {N-[2,4-dimethyl-5-[[(trifluoromethyl)sulfonyl]amino]phenyl]acetamide} were usually ineffective. Clopyralid + picloram, triclopyr, or 2,4,5-T applied in 1:1 combinations at 0.07 + 0.07 kg/ha were usually equally or more effective than any one of the herbicides applied alone at 0.14 kg/ha. Adjuvants did not enhance the phytotoxicity of picloram, triclopyr, or 2,4,5-T on field-grown honey mesquite but sometimes did with clopyralid.     

Preference for feeding on honey solution and its effect on survival,development, and fecundity of Ebrechtella tricuspidata

Members of the spider families Thomisidae, Salticidae, Miturgidae, Anyphaenidae, and Corinnidae have been observed on floral and extrafloral nectaries of plants, where they are presumably feeding on nectar. Nectar feeding is a phenomenon that has long intrigued biologists. However, few studies have focused on the effect of nectar on biological characteristics of these spiders. The aim of this study was to determine whether crab spiders, Ebrechtella tricuspidata (Fabricius) (Araneae: Thomisidae), show an active preference for honey solution (a simulated nectar source) and to test the effect of honey solution on the survival and development time of spiderlings and the fecundity of adult E. tricuspidata. The results showed that the number of females feeding on the 10, 20, and 30% honey solution was significantly higher than those feeding on water alone, and there was no difference in the number of females feeding on the three concentrations of honey solution. Significantly more E. tricuspidata preferred feeding on the 20% honey solution, and they spent significantly more time feeding on 20% honey solution than on water, regardless of whether they were males or females, adults or spiderlings. Ingestion of honey solution significantly increased the survival and shortened the development time of E. tricuspidata. Female spiders that fed on honey solution had a shorter pre‐oviposition period and laid more eggs than those given only water. Our results suggest that nectar could be a high‐quality supplementary food to maintain normal growth and metabolism in spiderlings and adult female spiders in nature.     

The amino acid composition of gum exudates from Prosopis species

The amino acid compositions of the proteinaceous components of the gum exudates from Prosopis alba, P. chilensis, P. glandulosa, P. laevigata, P. torreyana and P. velutina, and for a sample of commercial gum mesquite, are presented. In agreement with data published previously for the polysaccharide components of their gums, only minor differences in composition are shown by these species. The amino acid compositions are characterized by very high proportions of hydroxyproline and by high proportions of proline and serine; these three amino acids account for 62.5% of those present in the gum from Prosopis velutina. The amino acid compositions of these Prosopis gums are remarkably similar to that established recently for the gum from Acacia senegal (gum arabic).