A facultative mutualism between aphids and an invasive ant increases plant reproduction

1. The consequences to plants of ant–aphid mutualisms, particularly those involving invasive ants, are poorly studied. Ant–aphid mutualisms may increase or decrease plant fitness depending on the relative cost of herbivory by ant‐tended aphids versus the relative benefit of increased ant suppression of other (non‐aphid) herbivores. 2. We conducted field and greenhouse experiments in which we manipulated the presence and absence of cotton aphids (Aphis gossypii) on cotton plants to test the hypothesis that a mutualism between cotton aphids and an invasive ant, the red imported fire ant (Solenopsis invicta), benefits cotton plants by increasing fire ant suppression of caterpillars. We also manipulated caterpillar abundance to test whether the benefit of the mutualism varied with caterpillar density. 3. We found that more fire ants foraged on plants with cotton aphids than on plants without cotton aphids, which resulted in a significant reduction in caterpillar survival and caterpillar herbivory of leaves, flower buds, and bolls on plants with aphids. Consequently, cotton aphids indirectly increased cotton reproduction: plants with cotton aphids produced 16% more bolls, 25% more seeds, and 10% greater seedcotton mass than plants without aphids. The indirect benefit of cotton aphids, however, varied with caterpillar density: the number of bolls per plant at harvest was 32% greater on plants with aphids than on plants without aphids at high caterpillar density, versus just 3% greater at low caterpillar density. 4. Our results highlight the potential benefit to plants that host ant–hemipteran mutualisms and provide the first experimental evidence that the consequences to plants of an ant–aphid mutualism vary at different densities of non‐aphid herbivores.     

Hormonal Regulation of Pedicel Abscission in Begonia Flower Buds

In order to analyse the hormonal regulation of flower bud shedding in Begonia, levels of indoleacetic acid (IAA), abscisic acid (ABA) and ethylene were determined in buds and pedicels. The translocation and metabolism of ¹⁴C-labeled IAA in pedicel segments were also studied. In a monoecious Begonia fuchsioides hybrid, abscising male flower buds contain about 1% of the IAA present in non-abscising female flowers. In a male Begonia davisii hybrid, the seasonal variation in bud drop coincides with changes in the IAA content of the buds, while also the release of IAA from the bud to the pedicel is hampered. Abscission zones of these pedicels always contain abscission promoting ethylene concentrations. The tissue is prevented from responding with abscission by IAA from the flower buds. The buds also contain ABA but without influencing abscission considerably. Pretreatment with ethylene or ABA does not affect IAA transport in pedicel segments. The rate of this transport is 4–6 mm × h^–1:; the capacity increases with the transverse area. In young segments, IAA is decarboxylated and also otherwise metabolized.     

Loss of flower bud vigour in the Mediterranean shrub,Cistus albidus L. at advanced developmental stages

To better understand aging in perennials, age‐related changes in the physiology of leaves and flower buds of the Mediterranean shrub, Cistus albidus L. were evaluated. Two groups of different ages (5 and 10 years old), both at advanced developmental stages but of similar size, were compared. Total plant biomass, biomass produced per apical meristem and levels of cytokinins, abscisic acid and jasmonic acid in leaves and flower buds, as well as flower production, were measured. No differences in plant size, vegetative growth rates and levels of phytohormones in leaves were observed between 5‐ and 10‐year‐old plants. However, they showed significant differences in flower bud development; the older plants having reduced vigour, with 29.6% of flowers reaching anthesis compared to 52.5% in the younger plants. Furthermore, endogenous concentrations of zeatin and abscisic acid in flower buds at stage I (start of flower organ formation) were 61% and 41%, respectively, smaller in 10‐ than in 5‐year‐old plants. At stage II (with all flower organs formed), zeatin and abscisic acid concentrations decreased by ca. 90% and 80%, respectively, but differences between age groups were still evident (60% and 29% for zeatin and abscisic acid, respectively). Jasmonic acid levels in flower buds decreased by 80% from stage I to II, but did not differ between age groups. Despite reductions in flower bud vigour, total number of flowers per individual was not significantly different between age groups, so that an age‐related loss in reproductive vigour at the organ level did not lead to a decrease in flower production at the whole plant level.     

Influence of intra-tree variation in time of budburst of white spruce on herbivory and the behaviour and survivorship of Zeiraphera canadensis

Abstract.

• 1 Field studies were carried out to determine the effects of intra-tree variation in the time of budburst of white spruce, Picea glauca Moench (Voss.), on the behaviour and survivorship of, and herbivory by, the spruce bud moth, Zeiraphera canadensis Mutt. & Free.
• 2 There was significant variation in the time of budburst among whorls, shoots and buds. Budburst was acropetal, with buds in the interior of the lower crown bursting first and terminal buds on terminal shoots in the upper crown bursting last.
• 3 Bud moths laid the greatest proportion of their eggs in the middle of the crown and egg hatch was usually best synchronized to budburst in this region. Many eggs hatched before terminal buds on terminal shoots in the upper crown had burst and thus intra-tree variation in budburst decreased the probability that first-instar larvae would colonize the most important plant parts for growth.
• 4 However, many later instars dispersed upwards and outwards in the crown and colonized the late bursting buds in the upper crown. Such dispersal reduces the effectiveness of intra-plant variation in budburst to reduce herbivory and permits Z.canadensis to eat young nutritious buds for a longer period of time.
• 5 Intra-tree variance in the date of budburst was greater than that between trees but there were no consistent differences between the intra-tree variance of trees in half-sib families with high or low susceptibility to Z.canadensis. Differences between trees in herbivory, bud moth density and survivorship were not related to the amount of intra-tree variation in budburst.

     

On the inability of ants to protect their plant partners and the effect of herbivores on different stages of plant reproduction

Ants co‐occur with herbivorous thrips in several tropical plants, but their interactions are largely unexplored. Should thrips be deterred by ants, a positive effect of ants on plant fitness might be expected. Here, by using an experimental study design with ant‐present and ant‐excluded treatments, we investigated the influence of Camponotus blandus on Pseudophilothrips obscuricornis abundance and herbivory in three extrafloral nectaried species: Banisteriopsis malifolia, B. laevifolia and B. stellaris. In addition, we examined the effect of thrips herbivory on flower set and fruit development and dispersion. Thrips abundance and herbivory were higher on ant‐present stems of B. malifolia and B. laevifolia, where thrips managed to escape from ants by hiding in between clusters of flower buds (thygmotaxis behaviour). In B. stellaris the results were the opposite, as flower bud clusters did not offer hiding places, so thrips were unable to hide from ants; thus both thrips abundance and herbivory were lower on ant‐present stems. Thrips herbivory had no significant effect on flower and fruit set, but samaras (V‐shaped winged fruits of Malpighiaceae) attacked by thrips presented severe distortions and asymmetries. This caused damaged fruits to be dispersed closer to the mother plant, whereas uninjured fruits were dispersed further away. This study is evidence that ant–plant–herbivore systems have variable outcomes depending on the species involved, their behaviour and the plant structure under consideration. Unlike other herbivores, thrips negatively influence the very last stage of plant reproduction. The minute and furtive herbivorous thrips have long been ignored in natural systems, but because of their wide host range, they may be important herbivores even in extrafloral nectaried plants, which are usually fiercely protected by ants.     

Con‐specific neighbours may enhance compensation capacity in an invasive plant

Facilitation, both by inter‐ and intra‐specific neighbours, is known to be an important process in structuring plant communities. However, only a small number of experiments have been reported on facilitation in plant invasions, especially between invasive con‐specific individuals. Here, we focus on how con‐specific neighbours of the invasive alien plant alligator weed affect the tolerance of alligator weed to herbivory by the introduced biological control agent, Agasicles hygrophila. We conducted greenhouse and garden experiments in which invasive plant density and herbivory intensity (artificial clipping and real herbivory) were manipulated. In the greenhouse experiment, artificial clipping significantly reduced plant biomass when plants were grown individually, but when con‐specific neighbours were present in the same pot, biomass was not significantly different from control plants. Similarly, when compared to control plants, plants that were subjected to herbivory by A. hygrophila produced more biomass when grown with two con‐specific neighbours than when grown alone. Real herbivory also resulted in an increased number of vegetative buds, and again when two con‐specific neighbours were present this effect was increased (a 55.3% increase in buds when there was no neighbour, but a 111.6% increase in buds when two con‐specific neighbours were present). In the garden experiment, in which plants were grown at high density (6 plants per pot), alligator weed fully recovered from defoliation caused by insects at levels from 20–30% to 100%. Our results indicate that the con‐specific association may increase the compensatory ability to cope with intense damage in this invasive plant.     

Latitudinal variation in suppression of flower bud formation at high temperature in four native dandelions (Taraxacum spp.) with different distributions and genetic structures in Japan

The control of the response of flowering to temperature plays a key role in successful range‐expansion of plants. A previous study showed that the suppression of flower‐bud formation at high temperature in Taraxacum officinale decreases genetically with latitude from north to south in Japan. The present study investigated whether similar trait variation occurs among populations of native Taraxacum species in Japan. Seedlings of T. albidum (a low‐ and mid‐latitude allopolyploid), T. japonicum (a mid‐latitude diploid) and T. venustum (a high‐latitude autopolyploid) were grown at three temperatures. Time to flower‐bud appearance increased with temperature in T. japonicum and T. venustum, but did not increase in T. albidum. Time to flower‐bud appearance did not differ significantly among the three species at 14°C, but it was shorter in T. albidum than in the other two species at 19°C and 24°C. The early appearance of buds of T. albidum was confirmed by another experiment in which plants of 18 populations from the three species and T. platycarpum (a mid‐latitude diploid) grown at 19°C were used. The results clearly indicate that high‐temperature suppression of flower‐bud formation was lower in low‐latitude species than in high‐latitude species. This interspecific variation is analogous to the intraspecific variation in T. officinale. Time to bud appearance of five populations in T. albidum was homogeneous within and between the populations. The results suggest that the five populations are monoclonal and lack the sensitivity of suppression of flower‐bud formation to high temperature.     

Physiologic and Yield Responses of Shaded Cotton to the Plant Growth Regulator PGR-IV

The plant growth regulator PGR-IV has been reported to improve the growth, boll retention, and yield of cotton (Gossypium hirsutum L.) under optimum growing conditions. However, little is known about the response of cotton to PGR-IV under low light stress. A 3-year field study was conducted to determine if applying PGR-IV before an 8-day period of shade (63% light reduction) benefitted the growth and yield of shaded cotton. Shading during early squaring did not affect yield. Shading after the first flower stage significantly increased leaf chlorophyll concentration and fruit abscission and decreased the leaf photosynthetic rate, nonstructural carbohydrate concentrations, and lint yield. Foliar application of PGR-IV at 292 mL ha⁻¹ at early squaring and first flower did not improve the leaf photosynthetic rate of shaded cotton. However, shaded plants receiving PGR-IV had higher nonstructural carbohydrate concentrations in the floral buds and significantly lower fruit abscission than the shaded plants without PGR-IV. Applying PGR-IV to the foliage before shading resulted in a numeric increase (6–18%) in lint yield compared with shaded plants without PGR-IV. The decreased fruit abscission from the application of PGR-IV was associated with improved assimilate translocation. The yield enhancement from foliar application of PGR-IV was attributed to increased fruit retention. However, the average boll weight of shaded plants with PGR-IV tended to be lower than that of shaded plants without PGR-IV. Lint percentage was not affected by PGR-IV. Foliar application of PGR-IV appears beneficial for increasing the fruit retention of shaded cotton. Received June 12, 1997; accepted January 19, 1998     

Inhibition of flower bud abscission and ethylene evolution by light and silver thiosulphate in Lilium

The ethylene evolution from 3 cm-long flower buds of Lilium X 'Enchantment' increased when the plants were placed in darkness. Also shortening the daily light period led to an increase in ethylene evolution from the buds. Dark-enhanced ethylene evolution could only be evoked in flower buds between 2 and 3.5 cm long. Flower bud abscission induced by dakrness could be prevented by injecting silver thiosulphate into the flower buds, or by using plants grown from bulbs previously immersed in a silver thiosulphate solution. It is suggested that light-controlled flower bud abscission in Lilium X 'Enchantment' is mediated by ethylene, and that an increase in ethylene evolution in darkness is a consequence of the influence of light on ethylene action rather than a direct influence on ethylene biosynthesis.     

Abscission of cotton flower buds and petioles caused by protein from boll weevil larvae

Protein extracted from boll weevil larvae causes abscission of cotton flower buds and debladed petioles. Abscission of buds is preceded by a rapid and persistent flaring of bracts; cell proliferation is induced in the abscission zone of debladed petioles. A possible mode of transmission of the agent from larva to bud tissue is discussed.     

Sexual differences in flower defense and correlated male‐biased florivory in a plant–florivore system

Male‐biased florivory is a prominent phenomenon in the interaction between plants and florivores, and is potentially related to the evolution of flower traits and sex expression; however, its adaptive significance is not well understood. We studied florivory in the geometrid moth Chloroclystis excisa utilizing flower buds of a sexually polymorphic shrub, Eurya japonica, to reveal relationships between flower sex and moth oviposition preference, moth larval performance, and plant phenolics composition. In the field, C. excisa exploited flower buds on male and hermaphrodite trees but never those on female trees. In the laboratory, moths showed a strong oviposition preference for male over female flower buds, and larvae did not survive on female buds. Mortality was caused solely by feeding on the calyx covering the female bud. Female calyces contained higher concentrations of total phenolics and condensed tannins than did male calyces. These results suggest that substantial sexual differences in defense against florivory may have evolved in association with the differentiation of flower sexes and that a strong preference for the weakly defended flower sex may have evolved in florivores as a counter‐adaptation.     

Plant glandular trichomes mediate protective mutualism in a spider–plant system

1. Although several species of Peucetia (Oxyopidae) live strictly in association with plants bearing glandular trichomes worldwide, to date little is known about whether these associations are mutualistic. 2. In this study we manipulated the presence of Peucetia flava on the glandular plant Rhynchanthera dichotoma in the rainy and post‐rain season, to test the strength of its effects on leaf, bud, and flower damage and plant reproductive output. In addition, we ran independent field experiments to examine whether these sticky structures improve spider fidelity to plants. 3. Peucetia suppressed some species of foliar phytophages, but not others. Although spiders have reduced levels of leaf herbivory, this phenomenon was temporally conditional, i.e. occurred only in the post‐rain but not in the rainy season. Floral herbivory was also reduced in the presence of spiders, but these predators did not affect plant fitness components. 4. Plants that had their glandular trichomes removed retained fewer insects than those bearing such structures. Spiders remained longer on plants with glandular trichomes than on plants in which these structures had been removed. Isotopic analyses showed that spiders that fed on live and dead labelled flies adhered to the glandular hairs in similar proportions. 5. Spiders incurred no costs to the plants, but can potentially increase individual plant fitness by reducing damage to reproductive tissues. Temporal conditionality probably occurred because plant productivity exceeded herbivore consumption, thus dampening top‐down effects. Specialisation to live on glandular plants may have favoured scavenging behaviour in Peucetia, possibly an adaptation to periods of food scarcity.     

Characterization of the damage of Spodoptera eridania (Cramer) and Spodoptera cosmioides (Walker) (Lepidoptera: Noctuidae) to structures of cotton plants

The cotton plant, Gossypium hirsutum, hosts various pests that damage different structures. Among these pests, Spodoptera cosmioides (Walker) and Spodoptera eridania (Cramer) (Lepidoptera: Noctuidae) are considered important. The objectives of this study were to characterize and to quantify the potential damage of S. eridania and S. cosmioides feeding on different structures of cotton plants. For this purpose, newly-hatched larvae were reared on the following plant parts: leaf and flower bud; leaf and boll; flower bud or boll; and leaf, flower bud and boll. The survival of S. cosmioides and S. eridania was greater than 80% and 70% for larvae fed on cotton plant parts offered separately or together, respectively. One larva of S. eridania damaged 1.7 flower buds, but did not damage bolls, while one larva of S. cosmioides damaged 5.2 flower buds and 3.0 cotton bolls. Spodoptera eridania and S. cosmioides can be considered species with potential to cause economic damage to cotton plants because they can occur throughout cotton developmental stages causing defoliation and losses of reproductive structures. Therefore, the results validate field observations that these two species of Spodoptera are potential pests for cotton.     

Heterogeneity in vertebrate and invertebrate herbivory and its consequences for New Zealand mistletoes

Vertebrate herbivores generally have greater effects than invertebrates on plants. However, few studies have investigated the effects of both invertebrate and vertebrate herbivores on a single plant species. In New Zealand, nationwide declines in mistletoe populations have often been attributed to possum herbivory, but never to insect herbivory. The main goal of the present study was to document levels of vertebrate and invertebrate herbivory on endemic New Zealand mistletoe plants to suggest whether herbivory is leading to mistletoe decline. In the present study, the annual amount of leaf loss from herbivory by the brushtail possum (Trichosurus vulpecula), insect herbivory and leaf abscission were measured in two populations each of three mistletoe species (Alepis flavida, Peraxilla colensoi, and Peraxilla tetrapetala, Loranthaceae). In two populations of each species from February 1997 to February 1998, abscission accounted for the most leaf loss (range 10–84% of total mean leaf area, mean 33%), whereas insects and possums usually removed small and similar amounts (less than 3%). Possum browse caused large amounts of abscission in only one population (A. flavida at Eglinton). Observed possum browse was more heterogeneous than insect browse among branches within a plant (possum coefficient of variation = 2.63, insect CV = 1.98, P < 0.001), among plants in a population (possum CV = 2.15, insect CV = 0.69, P < 0.001), and between populations (possum CV = 1.36, insect CV = 1.09). Moreover, insects damaged 100% of the study plants but never removed more than 16% of leaf area on a single plant, whereas possums only browsed 32% of the study plants but severely defoliated some plants. Thus, while the mean amount of biomass removed across a population may have important consequences for mistletoe survival, the effect of possums on mistletoe populations may also depend on the heterogeneity of browse among individuals in the population.     

Floral herbivory at different stages of flower development changes reproduction in <Emphasis Type="Italic">Iris gracilipes</Emphasis> (Iridaceae)

Floral herbivores and pollinators are major determinants of plant reproduction. Because interaction of floral herbivores and pollinators occurs when herbivores attack the flowers in the bud and flower stages and because the compensatory ability of plants is known to differ according to the timing of herbivory, the effects of herbivory may differ according to its timing. In this study, we investigated the effects of floral herbivory at different stages on fruit production and seed/ovule ratio at two sites of large populations of the perennial herb, Iris gracilipes for 2 years. Herbivory at the bud and fruit stages both tended to have negative effects on fruit production, but the former caused more severe damage. On the other hand, herbivory at the flower stage tended not to have negative effects on fruit production because the degree of flower loss was smaller in the flower stage. Although herbivory decreased fruit production, flowers did not compensate for the damage by increasing the seed/ovule ratio because reproduction of I. gracilipes was limited by pollen availability rather than resources. These results indicate that floral herbivory at different stages has different effects on plant reproduction.     

Compensation to simulated insect leaf herbivory in wild cotton (Gossypium hirsutum): responses to multiple levels of damage and associated traits

• Identifying the mechanisms of compensation to insect herbivory remains a major challenge in plant biology and evolutionary ecology. Most previous studies have addressed plant compensatory responses to one or two levels of insect herbivory, and the underlying traits mediating such responses remain elusive in many cases.
• We evaluated responses associated with compensation to multiple intensities of leaf damage (0% control, 10%, 25%, 50%, 75% of leaf area removed) by means of mechanical removal of foliar tissue and application of a caterpillar (Spodoptera exigua) oral secretions in 3‐month‐old wild cotton plants (Gossypium hirsutum). Four weeks post‐treatment, we measured plant growth and multiple traits associated with compensation, namely: changes in above‐ and belowground, biomass and the concentration of nutrients (nitrogen and phosphorus) and non‐structural carbon reserves (starch and soluble sugars) in roots, stems and leaves.
• We found that wild cotton fully compensated in terms of growth and biomass allocation when leaf damage was low (10%), whereas moderate (25%) to high leaf damage in some cases led to under‐compensation. Nonetheless, high levels of leaf removal (50% and 75%) in most cases did not cause further reductions in height and allocation to leaf and stem biomass relative to low and moderate damage. There were significant positive effects of leaf damage on P concentration in leaves and stems, but not roots, as well as a negative effect on soluble sugars in roots.
• These results indicate that wild cotton fully compensated for a low level of leaf damage but under‐compensated under moderate to high leaf damage, but can nonetheless sustain growth despite increasing losses to herbivory. Such responses were possibly mediated by a re‐allocation of carbohydrate reserves from roots to shoots.

     

Olfactory response of a predatory mirid to herbivore induced plant volatiles: multiple herbivory vs. single herbivory

Plants infested with a single herbivore species can attract natural enemies through the emission of herbivore‐induced plant volatiles (HIPVs). However, under natural conditions plants are often attacked by more than one herbivore species. We investigated the olfactory response of a generalist predators Macrolophus caliginosus to pepper infested with two‐spotted spider mites, Tetranychus urticae, or green peach aphid, Myzus persicae, vs. plants infested with both herbivore species in a Y‐tube olfactometer set up. In addition, the constituents of volatile blends from plants exposed to multiple or single herbivory were identified by gas chromatography‐mass spectrometry (GC‐MS). The mirid bugs showed a stronger response to volatiles emitted from plants simultaneously infested with spider mites and aphids than to those emitted from plants infested by just one herbivore, irrespective of the species. Combined with results from previous studies under similar conditions we infer that this was a reaction to herbivore induced plant volatiles. The GC‐MS analysis showed that single herbivory induced the release of 22 additional compounds as compared with the volatiles emitted from clean plants. Quantitative analyses revealed that the amount of volatile blends emitted from pepper infested by both herbivores was significantly higher than that from pepper infested by a single herbivore. Moreover, two unique substances were tentatively identified (with a probability of 94% and 91%, respectively) in volatiles emitted by multiple herbivory damaged plants: α‐zingiberene and dodecyl acetate.     

Studies on the abscission of flowers and fruits of cotton (Gossypium barbadense L.)

Effects of exogenous application of auxin, GA3, abscisic acid, ethrel, methionine and α-alanine to the cut ends of the pedicels of flower buds, flowers and fruits on their abscission behaviour were studied. Fruit pedicels required more time for abscission compared with flower and flower bud pedicels. NAA inhibited abscission of all types of pedicels and the inhibition was maximum in matured fruit pedicels and minimum in flower bud pedicels. Flower pedicels were more sensitive towards the abscission promotive effects of GA3, abscisic acid and ±-alanine and the flower bud pedicels towards ethrel and methionine. The duration of Stage-I of abscission was maximum in cut pedicels of fruit and minimum in those of flower buds. Biochemical analyses revealed greater quantities of endogenous amino acids in the epicalyx of flowers with the exception of methionine and aspartic acid which were found to be present in higher quantities in the epicalyx of flower buds. Levels of IAA-like compounds were maximum in the epioalyx of flower buds and minimum in the epicalyx of flowers. Higher levels of abscisic acid were found in the epicalyx of matured fruits and the epicalyx of flower buds showed a minimum amount of abscisic acid-like compound.