Stem respiration of ponderosa pines grown in contrasting climates: implications for global climate change

We examined the effects of climate and allocation patterns on stem respiration in ponderosa pine (Pinus ponderosa) growing on identical substrate in the cool, moist Sierra Nevada mountains and the warm, dry, Great Basin Desert. These environments are representative of current climatic conditions and those predicted to accompany a doubling of atmospheric CO₂, respectively, throughout the range of many western north American conifers. A previous study found that trees growing in the desert allocate proportionally more biomass to sapwood and less to leaf area than montane trees. We tested the hypothesis that respiration rates of sapwood are lower in desert trees than in montane trees due to reduced stem maintenance respiration (physiological acclimation) or reduced construction cost of stem tissue (structural acclimation). Maintenance respiration per unit sapwood volume at 15°C did not differ between populations (desert: 6.39 ± 1.14 SE μmol m⁻³ s⁻¹, montane: 6.54 ± 1.13 SE μmol m⁻³ s⁻¹, P = 0.71) and declined with increasing stem diameter (P = 0.001). The temperature coefficient of respiration (Q ₁₀) varied seasonally within both environments (P = 0.05). Construction cost of stem sapwood was the same in both environments (desert: 1.46 ± 0.009 SE g glucose g⁻¹ sapwood, montane: 1.48 ± 0.009 SE glucose g⁻¹ sapwood, P = 0.14). Annual construction respiration calculated from construction cost, percent carbon and relative growth rate was greater in montane populations due to higher growth rates. These data provide no evidence of respiratory acclimation by desert trees. Estimated yearly stem maintenance respiration was greater in large desert trees than in large montane trees because of higher temperatures in the desert and because of increased allocation of biomass to sapwood. By analogy, these data suggest that under predicted increases in temperature and aridity, potential increases in aboveground carbon gain due to enhanced photosynthetic rates may be partially offset by increases in maintenance respiration in large trees growing in CO₂-enriched atmospheres. Received: 4 November 1996 / Accepted: 23 January 1997     

Wound-inducible and organ-specific expression of ORF13 from Agrobacterium rhizogenes 8196 T-DNA in transgenic tobacco plants

Tissue-specific expression of the ORF13 promoter from Agrobacterium rhizogenes 8196 was assessed throughout the development of transgenic tobacco plants using a GUS reporter gene. ORF13 exhibited high activity in roots but with different patterns of expression. The activity of the ORF13 promoter in vascular tissues increased from the base to the tip of the stem. The ORF13 promoter is wound inducible in a limited area adjacent to the wound site. The time course of wound induction of ORF13 in transgenic tobacco containing an ORF13 promoter-GUS translational fusion was similar to that previously described for genes involved in plant defense responses. A series of 5′ deletions of the ORF13 promoter fused to the β-glucuronidase gene was examined for expression in roots and leaves of transgenic plants. Cis-acting elements that modulate quantitative expression of the transgene after wounding were detected. Received: 11 July 1996 / Accepted: 19 November 1996     

Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success. Received: 21 March 1996 / Accepted: 24 October 1996     

Odour-mediated responses of phytophagous mites to conspecific and heterospecific competitors

Plants under herbivore attack produce volatiles, thus attracting natural enemies of the herbivores. However, in doing so, the plant becomes more conspicuous to other herbivores. Herbivores may use the odours as a cue to refrain from visiting plants that are already infested, thereby avoiding competition for food, or, alternatively, to visit plants with defences weakened by earlier attacks. We investigated the response of one species of herbivore (the spider mite Tetranychus urticae) to odours emanating from cucumber plants infested by conspecific or heterospecific (the western flower thrips, Frankliniella occidentalis) herbivores. Olfactometer experiments in the laboratory showed that spider mites have a slight, but significant, preference for plants infested with conspecifics, but strongly avoid plants with thrips. These results were substantiated with greenhouse experiments. We released spider mites on the soil in the centre of a circle of six plants, half of which were infested with either conspecifics or heterospecifics (thrips), whereas the other half were uninfested. It was found that 60–70% of the mites were recaptured on the plants within 5 h after release. Results of these experiments were in agreement with results of the olfactometer experiments: (1) significantly fewer spider mites were found on plants infested with thrips than on uninfested plants and (2) more mites were found on plants with conspecifics than on clean plants (although this difference was not significant). From a functional point of view it makes sense that spider mites prefer clean plants over thrips-infested plants, since thrips are not only competitors, but are also known as intraguild predators of spider mites. Possible reasons for the slight attraction of spider mites to plants infested with conspecifics are discussed. Received: 22 June 1996 / Accepted: 29 September 1996     

Effects of plant hybridization on herbivore-parasitoid interactions

We studied the effects of host plant hybridization on the survival and mortality of the leaf-mining moth Phyllonorycter salicifoliella on hybrid and parental willow plants in the field and in a common garden experiment. P. salicifoliella survival differed significantly among three willow taxa in the field in 1994 but not in the field in 1995 or in the common garden. Parasitism by eulophid wasps differed significantly among taxa in 1994 and appeared to account for the variation in their survival. In the field in 1995, host feeding predation varied significant among taxa. The theory of tritrophic interactions predicts that plant genotype can affect natural enemy impact, and this study supports this prediction. Significant variation in survival and eulophid parasitism was also found among genotypes within taxa in the field in both years and in the common garden experiment. The common garden results show that genetic differences in plants affect the herbivore-parasitoid interaction. Variation among years in the patterns of survival and causes of mortality among field plants suggest that genotype by environment interactions may be important. Received: 1 March 1996 / Accepted: 4 November 1996     

Body fluid homeostasis and cardiovascular adjustments during submaximal exercise: influence of chewing coca leaves

 The present study was undertaken to determine the haematological and cardiovascular status, at rest and during prolonged (1 h) submaximal exercise (approximately 70% of peak oxygen uptake) in a group (n = 12) of chronic coca users after chewing approximately 50 g of coca leaves. The results were compared to those obtained in a group (n = 12) of nonchewers. At rest, coca chewing was accompanied by a significant increase in heart rate [from 60 (SEM 4) TO 76 (SEM 3) beats · min⁻¹], in haematocrit [from 53.2 (SEM 1.2) to 55.6 (SEM 1.1)%] in haemoglobin concentration, and plasma noradrenaline concentration [from 2.8 (SEM 0.4) to 5.0 (SEM 0.5) μmol · l⁻¹]. It was calculated that coca chewing for 1 h resulted in a significant decrease in blood [−4.3 (SEM 2.2)%] and plasma [−8.7 (SEM 1.2)%] volume. During submaximal exercise, coca chewers displayed a significantly higher heart rate and mean arterial blood pressure. The exercise-induced haemoconcentration was blunted in coca chewers compared to nonchewers. It was concluded that the coca-induced fluid shift observed at rest in these coca chewers was not cumulative with that of exercise, and that the hypovolaemia induced by coca chewing at rest compromised circulatory adjustments during exercise. Accepted: 29 October 1996     

Discrimination of sinusoidally frequency-modulated sound signals mimicking species-specific communication calls in the FM-bat Phyllostomus discolor

In the lesser spear-nosed bat, Phyllostomus discolor, maternal directive calls are characterized by an individual type of sinusoidal frequency modulation (= SFM) pattern. Beside modulation frequency, modulation depth, carrier frequency, and number of modulation cycles per call contribute to the mother's vocal signature. Since juvenile P. discolor learn to adapt their isolation calls to the corresponding call characteristics of the own mother or even to playback of a computer-stored directive call, if hand-reared in the absence of conspecifics, the bats' auditory system ought to be able to resolve interindividual differences in communication call structure. However, quantitative psychoacoustic data on the discrimination of SFM signals in this species are not available. Thus, in the present study, lesser spear-nosed bats were trained in a two-alternative forced-choice procedure to discriminate between two alternatingly presented SFM sound signals differing in modulation frequency. Other characteristics of acoustic stimuli were identical and designed to mimick the fundamental of species-specific calls. By gradually reducing the difference in modulation frequency between both stimuli within the behavioural relevant range until the animals' performance dropped below the 75%-correct level, a considerable auditory spectro-temporal resolution has been revealed. Particularly in comparison to the overall interindividual variation of this call parameter (minimal modulation frequency = 49 Hz, maximum = 100 Hz), the determined average difference limen for modulation frequency of 2.42 ± 0.29 Hz seems substantial and sufficient for labelling individuals. Accepted: 30 November 1996     

High vapour pressure deficit exacerbates xylem cavitation and photoinhibition in shade-grown Piper auritum H.B. & K. during prolonged sunflecks

Water relations dynamics during simulated sunflecks at high (36°C) and medium (27°C) temperatures and high and low vapour pressure deficits beween leaf and air (VPD) were studied on shade-grown Piper auritum H.B. & K. plants, a pioneer tree, common in gaps and clearings of tropical rain forests. The leaves of P. auritum wilt rapidly when exposed to high light. Exposure to high VPD and high light caused substantial and rapid dehydration of leaves. Dehydration could be prevented under high humidity irrespective of temperature. Water stored in leaf cells served as initial source for transpiration upon high light exposure. This effect increased with increasing VPD and temperature. The pronounced decrease in leaf water content over time in high light caused a rapid decrease in leaf water potential (Ψ_l) and a concomitant increase in water potential gradient (ΔΨ/Δx) between trunk and leaf, yet the high leaf elasticity (small bulk elastic modulus, ε) allowed turgor maintenance under most conditions. Under high VPD and high temperature, stomata remained open and ΔΨ/Δx frequently exceeded 0.95 MPa · m⁻¹, the cavitation-inducing threshold (ΔΨ/Δx _cav) causing high rates of acoustic emissions from stems and leaf petioles and leading to concomitant losses in hydraulic conductance per leaf area (k _l). At medium temperature (high VPD), stomatal closure contained xylem embolism by keeping ΔΨ/Δx at or below this threshold. We argue that wilting substantially contributes to creating a sufficient driving force for water uptake from the soil, and reducing the VPD (through a decrease in radiation load and thus leaf temperature) to avoid excessive dehydration. Received: 3 March 1996 / Accepted: 10 November 1996     

Blood flow in the carotid artery during breath-holding in relation to diving bradycardia

The present study investigated the mechanism of diving bradycardia. A group of 14 healthy untrained male subjects were examined during breath-holding either out of the water (30–33°C), in head-out immersion, or in whole-body submersion (27–29°C) in a diving pool. Blood velocity, blood volume flow in the carotid artery, diastolic blood pressure and electrocardiogram were measured and recorded during the experiments. The peak blood velocity increased by 13.6% (P < 0.01) and R-wave amplitude increased by 57.1% (P < 0.005) when the subjects entered water from air. End-diastolic blood velocity in the carotid artery increased significantly during breath-holding, e.g. increased from 0.20 (SD 0.02) m · s⁻¹ at rest to 0.33 (SD 0.04) m · s⁻¹ (P < 0.001) at 50.0 s in breath-hold submersion to a 2.0-m depth. Blood volume flow in the carotid artery increased by 26.6% (P < 0.05) at 30 s and 36.6% (P < 0.001) at 40 s in breath-hold submersion to a 2.0-m depth. Diastolic blood pressure increased by 15.4% (P < 0.01) at 60 s during breath-holding in head-out immersion. Blood volume flow, and diastolic blood pressure increased significantly more and faster during breath-holding in submersion than out of the water. There was a good negative correlation with the heart rate: the root mean square correlation coefficient r was 0.73 (P < 0.001). It was concluded that an increased accumulation of blood in the aorta and arteries at end-diastole and decreased venous return, caused by an increase in systemic peripheral resistance during breath-holding, underlies diving bradycardia. Accepted: 22 November 1996     

Effects of 3 days of carbohydrate supplementation on muscle glycogen content and utilisation during a 1-h cycling performance

This study compared the effects of supplementing the normal diets of six trained cyclists [maximal oxygen uptake O_2max) 4.5 (0.36)l · min⁻¹; values are mean (SD)] with additional carbohydrate (CHO) on muscle glycogen utilisation during a 1-h cycle time-trial (TT). Using a randomised crossover design, subjects consumed either their normal diet (NORM) for 3 days, which consisted of 426 (137) g · day⁻¹ CHO [5.9 (1.4) g · kg⁻¹ body mass (BM)], or additional CHO (SUPP) to increase their intake to 661 (76) g · day⁻¹ [9.3 (0.7) g · kg⁻¹ BM]. The SUPP diet elevated muscle glycogen content from 459 (83) to 565 (62) mmol · kg⁻¹ dry weight (d.w.) (P < 0.05). However, despite the increased pre-exercise muscle glycogen stores, there was no difference in the distance cycled during the TT [40.41 (1.44) vs 40.18 (1.76) km for NORM and SUPP, respectively]. With NORM, muscle glycogen declined from 459 (83) to 175 (64) mmol · kg⁻¹ d.w., whereas with SUPP the corresponding values were 565 (62) and 292 (113) mmol · kg⁻¹ d.w. Accordingly, both muscle glycogen utilisation [277 (64) vs 273 (114) mmol · kg⁻¹ d.w.] and total CHO oxidation [169 (20) vs 165 (30) g · h⁻¹ for NORM and SUPP, respectively] were similar. Neither were there any differences in plasma glucose or lactate concentrations during the two experimental trials. Plasma glucose concentration averaged 5.5 (0.5) and 5.6 (0.6) mmol · l⁻¹, while plasma lactate concentration averaged 4.4 (1.9) and 4.4 (2.3) mmol · l⁻¹ for NORM and SUPP, respectively. The results of this study show that when well-trained subjects increase the CHO content of their diet for 3 days from 6 to 9 g · kg⁻¹ BM there is only a modest increase in muscle glycogen content. Since supplementary CHO did not improve TT performance, we conclude that additional CHO provides no benefit to performance for athletes who compete in intense, continuous events lasting 1 h. Furthermore, the substantial muscle CHO reserves observed at the termination of exercise indicate that whole-muscle glycogen depletion does not determine fatigue at this exercise intensity and duration. Accepted: 25 November 1996     

The effect of a thiamin derivative on exercise performance

The purpose of this study was to investigate the effect of a thiamin derivative, thiamin tetrahydrofurfuryl disulfide (TTFD), on oxygen uptake (˙VO₂), lactate accumulation and cycling performance during exercise to exhaustion. Using a randomized, double-blind, cross-over design with a 10-day washout between trials, 14 subjects ingested either 1 g · day⁻¹ of TTFD or a placebo (PL) for 4 days. On day 3, subjects performed a progressive exercise test to exhaustion on a cycle ergometer for the determination of ˙VO_2submax, ˙VO_2peak, lactate concentration ([La⁻ ]), lactate threshold (Th_La) and heart rate ( f _c). On day 4, subjects performed a maximal 2000-m time trial on a cycle ergometer. A one-way analysis of variance (ANOVA) with repeated measures was used to determine significant differences between trials. There were no significant differences detected between trials for serial measures of ˙VO_2submax, [La⁻] or f _c. Likewise, ˙VO_2peak [PL 4.06 (0.19) TTFD 4.12 (0.19) l · min⁻¹, P = 0.83], Th_La [PL 2.47 (0.17), TTFD 2.43 (0.16) l · min⁻¹, P = 0.86] and 2000-m performance time [PL 204.5 (5.5), TTFD 200.9 (4.3) s, P = 0.61] were not significantly different between trials. The results of this study suggest that thiamin derivative supplementation does not influence high-intensity exercise performance. Accepted: 19 December 1996     

In vivo expression of a full-length cDNA copy of potato leafroll virus (PLRV) in protoplasts and transgenic plants

A full-length cDNA copy (PLRV_fl) of potato leafroll virus (PLRV) was constructed and examined in vivo for its biological activities by transient expression experiments with plasmid DNA or in vitro transcribed RNA. In addition, PLRV_fl cDNA was stably introduced into the genome of potato plants by Agrobacterium-mediated leaf disc transformation. Both transient and stable expression of PLRV_fl resulted in the synthesis of genomic and subgenomic PLRV RNAs. Transgenic plants accumulated the 17-kDa movement protein and displayed the typical symptoms of PLRV infection. This is the first example of the constitutive expression of a phloem-limited virus in planta. Received: 20 August 1996 / Accepted: 24 September 1996     

Rhythmic components of photoperiodic time measurement in the pitcher-plant mosquito, Wyeomyia smithii

Photoperiodic time measurement regulating larval diapause in the pitcher-plant mosquito, Wyeomyia smithii, varies in a close relationship with latitude. The critical photoperiod mediating the maintenance and termination of diapause is positively correlated with latitude (r ² = 0.977) among six populations from southern (30–31° N), intermediate (40° N), and northern (46–49° N) latitudes in North America. The developmental response to unnaturally short and to unnaturally long photoperiods declines with increasing latitude, so that longer critical photoperiods are associated with a downward rather than a lateral shift in the photoperiodic response curve. Exotic light and dark cycles of varying period (T) with a short (10 h) photophase and a scotophase ranging from 14 (T = 24) to 62 (T = 72) h, reveal two geographic patterns: a decline in perturbability of the photoperiodic clock with increasing latitude, and no change with latitude in the 21-h period of rising and falling development with increasing T. These results show (1) that there is a rhythmic component to photoperiodic time measurement in W. smithii, (2) that the period of this rhythm is about 21 h in all populations, and (3) that more northern populations show decreasing responsiveness to photoperiod and increasing stability against perturbation by exotic period lengths (T > 24). Previous studies on W.␣smithii indicate that this single temperate species of a tropical and subtropical genus has evolved from south to north. We therefore conclude that the evolution of increasing critical photoperiod in W. smithii during its adaptive radiation into North America has more likely involved the amplitude and not the period of the underlying circadian pacemaker. Received: 22 July 1996 / Accepted: 30 September 1996     

Populus tremuloides photosynthesis and crown architecture in response to elevated CO2 and soil N availability

We tested the hypothesis that elevated CO₂ would stimulate proportionally higher photosynthesis in the lower crown of Populus trees due to less N retranslocation, compared to tree crowns in ambient CO₂. Such a response could increase belowground C allocation, particularly in trees with an indeterminate growth pattern such as Populus tremuloides. Rooted cuttings of P. tremuloides were grown in ambient and twice ambient (elevated) CO₂ and in low and high soil N availability (89 ± 7 and 333 ± 16 ng N g⁻¹ day⁻¹ net mineralization, respectively) for 95 days using open-top chambers and open-bottom root boxes. Elevated CO₂ resulted in significantly higher maximum leaf photosynthesis (A _max) at both soil N levels. A _max was higher at high N than at low N soil in elevated, but not ambient CO₂. Photosynthetic N use efficiency was higher at elevated than ambient CO₂ in both soil types. Elevated CO₂ resulted in proportionally higher whole leaf A in the lower three-quarters to one-half of the crown for both soil types. At elevated CO₂ and high N availability, lower crown leaves had significantly lower ratios of carboxylation capacity to electron transport capacity (V _cmax/J _max) than at ambient CO₂ and/or low N availability. From the top to the bottom of the tree crowns, V _cmax/J _max increased in ambient CO₂, but it decreased in elevated CO₂ indicating a greater relative investment of N into light harvesting for the lower crown. Only the mid-crown leaves at both N levels exhibited photosynthetic down regulation to elevated CO₂. Stem biomass segments (consisting of three nodes and internodes) were compared to the total A _leaf for each segment. This analysis indicated that increased A _leaf at elevated CO₂ did not result in a proportional increase in local stem segment mass, suggesting that C allocation to sinks other than the local stem segment increased disproportionally. Since C allocated to roots in young Populus trees is primarily assimilated by leaves in the lower crown, the results of this study suggest a mechanism by which C allocation to roots in young trees may increase in elevated CO₂. Received: 12 August 1996 / Accepted: 12 November 1996     

c-fos expression in the putative avian suprachiasmatic nucleus

c-fos induction was investigated as a potential component in the avian photic entrainment pathway and as a possible means of locating the central pacemaker in birds. In both quail (Coturnix coturnix japonica) and starlings (Sturnus vulgaris) exposure to 1 h of light induced Fos-lir in the visual suprachiasmatic nucleus but not in the medial suprachiasmatic nucleus. However, the degree of c-fos induction in the visual suprachiasmatic nucleus was similar at different circadian times despite the fact that the light pulses caused differential phase shifts in the locomotor rhythm. For golden hamsters the same experiment resulted in significantly different levels of Fos-lir in the suprachiasmatic nucleus, as well as different phase shifts. Starlings and hamsters were also entrained to T-cycles that caused a large daily phase shift (T = 21.5 h in starlings, T = 22.67 hours in hamsters), or no daily phase shift (T = free running period). No difference in the induced levels of Fos-lir in the visual suprachiasmatic nucleus region was observed between the two groups of starlings, but in hamsters there were significantly different levels of Fos-lir in the suprachiasmatic nucleus between the two groups. Accepted: 15 November 1996     

Effect of masking the parietal eye on the diurnal activity and body temperature of two sympatric species of monitor lizards, Varanus s. salvator and Varanus b. nebulosus

 Five water monitor lizards, Varanus salvator salvator, and four clouded monitor lizards, Varanus bengalensis nebulosus, were caught on Tioman island in Malaysia. A radio-thermistor transmitter was implanted into the buccal cavity of each animal, and they were released into an enclosure measuring 5.5 × 6.5 metres. The lizards were observed for 9 and 8 days, respectively, before and after the parietal eye was covered with aluminium foil. With uncovered parietal eye, both species showed a clear diurnal rhythm, being active only during day time. After covering the parietal eye, the mean locomotor activity of five V. s. salvator decreased from 791 to 107 min · day^–1 but remained unchanged around 850 min · day^–1 for V. b. nebulosus. The mean duration of locomotor activity decreased in V. s. salvator and V. b. nebulosus after the parietal eye was covered, but V. b. nebulosus maintained its locomotor activity by increasing the number of locomotor bouts. The water monitor spent very little time on thermoregulation. Its body temperature ranged between 26.3 and 28.4 °C, which decreased after the parietal eye was covered. The clouded monitor thermoregulated around 28.8–36.0 °C, which remained unchanged after the parietal eye was covered. In both species, there was a strong correlation between body temperature and ambient temperature. Behavioural abnormalities were recorded among V. s. salvator with covered parietal eye. They were often observed to be active by night and often slept outside a burrow. The circadian rhythm of V. b. nebulosus appeared unaffected by shielding of its parietal eye. Captivity combined with shielded parietal eye induced agonistic behaviour in both species. Accepted: 11 September 1996     

Regulatory elements in the promoter of the vitelline membrane gene VM32E of Drosophila melanogaster direct gene expression in distinct domains of the follicular epithelium

The Drosophila vitelline membrane protein gene VM32E is expressed according to a precise temporal and spatial program in the follicle cells. Results from germ line transformation experiments using different fragments of the −465/−39 VM32E region fused to the hsp/lacZ reporter gene revealed that the region −348/−39 is sufficient to confer the wild-type expression pattern. Within this segment, distinct cis-regulatory elements control VM32E expression in ventral and dorsal follicle cells. The region between −135/−113 is essential for expression of the VM32E gene in the ventral columnar follicle cells. Expression in the dorsal domain requires the two regions −348/−254 and −118/−39. Furthermore, the region −253/−119 appears to contain a negative element that represses gene activity in anterior centripetal cells. We suggest that the expression of the VM32E gene throughout the follicular epithelium is controlled by specific cis-regulatory elements acting in distinct spatial domains and following a precise developmental program. Received: 22 October 1996 / Accepted: 14 November 1996     

Acorn selection by Mexican jays: a test of a tri-trophic symbiotic relationship hypothesis

By caching acorns, jays serve as important dispersal agents for oak (Quercus) species. Yet little is known about which acorn characteristics affect selection by jays. In the traditional model of jay/oak symbiosis, large, brown, ripe acorns free of invertebrate parasites (e.g., Curculio acorn weevils) are selected by jays. Recently, it has been suggested that a tri-trophic relationship between oaks, jays, and weevils may have evolved to counter the negative dietary effects of acorn tannins. Under the tri-trophic model, jays would preferentially select acorns containing weevil larvae. We tested the assumptions that (1) acorns containing curculionid larvae exist in sufficient quantities to support jay populations and (2) jays can detect, and preferentially select, acorns containing weevil larvae, and investigated the cues by which jays select acorns. Captive Mexican jays (Aphelocomaultramarina) were presented Emory oak (Quercusemoryi) acorns in aviary feeding trials. Large, dense, viable acorns free of curculionid larvae were preferentially selected. Contrary to results of previous research, color did not affect selection. Acorn viability increased and curculionid larval occupancy decreased in adjacent savannas and isolated stands relative to existing oak woodland, perhaps favoring oak recruitment into adjacent lower-elevation grasslands. Our results compel us to reject the tri-trophic model for this system, and are consistent with the traditional jay/oak symbiosis model. Relatively long-distance dispersal of viable acorns favors Emory oak replacement, and spatial patterns of acorn viability and curculionid parasitism suggest expansion of Emory oak into adjacent low-elevation semi-arid grasslands. Received: 29 February 1996 / Accepted: 26 September 1996