Allocation of nitrogen to an inducible defense and seed production in Nicotiana attenuata

Resource-based tradeoffs in the allocation of a limiting resource are commonly invoked to explain negative correlations between growth and defense in plants, but critical examinations of these tradeoffs are lacking. To rigorously quantify tradeoffs in a common currency, we grew Nicotiana attenuata plants in individual hydroponic chambers, induced nicotine production by treating roots with methyl jasmonate (MJ) and standardized leaf puncturing, and used ¹⁵N to determine whether nitrogen-based tradeoffs among nicotine production, growth, and seed production could be detected. Plants were treated with a range of MJ quantities (5, 45 or 250 μg plant^?1) to effect a physiologically realistic range of changes in endogenous jasmonic acid levels and increases in nicotine production and accumulation; MJ treatments were applied to the roots to target JA-induced nicotine production, since nicotine biosynthesis is restricted to the roots. Leaf puncturing and 5 μg MJ treatments increased de novo nicotine synthesis and whole-plant (WP) nicotine pools by 93 and 66%, while 250 μg MJ treatments increased these values 3.1 and 2.5-fold. At these high rates of nicotine production, plants incorporated 5.7% of current nitrogen uptake and 6.0% of their WP nitrogen pools into nicotine. The ¹⁵N-labeled nicotine pools were stable or increased for the duration of vegetative growth, indicating that the N-nicotine was not metabolized and re-used for growth. Plants with elevated nicotine production grew more slowly and the differences in plant biomass gain between MJ-treated plants and controls were linearly related to the differences in nicotine accumulation. Despite the reductions in rosette-stage growth associated with nicotine production, estimates of lifetime fitness (cumulative lifetime seed production, mass/seed, seed viability) were not affected by any treatment. Only two treatments (leaf puncturing and 250 μg MJ) increased the allocations of ¹⁵N acquired at the time of induction to seed production. On average, plants used only 14.9% of their WP nitrogen pool for seed production, indicating that either the nitrogen requirements for seed production or the reproductive effort of these hydroponically-grown plants are low. To determine if seed production is strongly influenced by the amount of vegetative biomass attained before reproduction, the experiment was repeated with plants that had 44% of their leaf area (or 29% of their WP biomass) removed before MJ treatments with a removal technique that minimized the nicotine response. MJ treatments of these plants dramatically increased nicotine production and accumulation, but these plants also suffered no measurable fitness consequences from either the leaf removal or MJ treatments. We conclude that when N. attenuata plants are grown in these individual hydroponic chambers, their allocation to reproduction is sufficiently buffered to obscure the large increases in nitrogen allocations to an inducible defense. To determine whether soil-grown plants are similarly buffered, we grew two genotypes of plants in the high-nutrient soil from a 1-year-old burn in a piñyon-juniper forest (the plants' natural habitat) and in low-nutrient soil from an adjacent unburned area, and induced nicotine production in half of the plants with a 500 μg root MJ treatment. Plants grown in burned soils had an estimated lifetime fitness that was on average 2.8-fold greater than that of plants grown in unburned soils. MJ treatment reduced fitness estimates by 43% and 71% in the burned and unburned soils, respectively. We conclude that while hydroponic culture allows one to rigorously quantitate nitrogen allocation to growth, reproduction and defense, the allocation patterns of plants grown in hydroponic culture differ from those of plants grown in soil. Under hydroponic conditions, plants have low reproductive allocations and reproductive-defense tradeoffs are not detected. Reproductive-defense tradeoffs are readily discernible in soil-grown plants, but under these growing conditions, the nitrogen-basis for the tradeoff is difficult to quantify.     

Induced responses in Nicotiana attenuata affect behavior and growth of the specialist herbivore Manduca sexta

Many plants employ induced responses against generalist herbivores. Specialist herbivores, however, may employ several mechanisms to overcome the negative effects of induced plant defenses. Here we test how the behavior and development of specialist Manduca sexta larvae are affected by induced responses in their natural host plant Nicotiana attenuata. On a spatial scale relevant to both the plant and the herbivore, we first determined how methyl jasmonate (MeJA)-induced responses, such as increased nicotine production, affect the tendency of larvae to leave induced plants. When larvae were allowed to move between two plants planted in one pot, they left an MeJA-treated plant faster than a control plant. When both plants in the pot were MeJA-treated, the larvae developed more slowly than when both plants were uninduced, or when the larvae had the opportunity to move to an uninduced neighbor. The sooner larvae moved from an MeJA-treated plant to an untreated neighbor, the larger the body mass they attained. This demonstrates that M. sexta larvae can compensate behaviorally for the deleterious effects of induced plant responses. These effects were observed in plants grown under both low and high N supply rates, though the effects were more pronounced under high N. To examine the consequences of the timing and the direction of the host plant switching behavior for larval development, neonate larvae were fed leaves excised from induced and uninduced plants. Larvae confined to MeJA-treated leaves had higher mortality rates and grew slower than larvae fed only control leaves. This demonstrates that MeJA-induced responses decrease growth and development of specialist herbivores that do not have the behavioral option of moving to an uninduced plant. The sooner the larvae were switched to MeJA-treated leaves, the slower their development compared to larvae fed only uninduced leaves. In contrast, the sooner larvae fed MeJA-treated leaves were switched to control leaves, the faster they developed. Again the effects of MeJA treatment were stronger in plants grown under high N supply. We propose that induced plants growing in close competition with an uninduced conspecific may offset the fitness costs of these induced responses and perhaps obtain a fitness benefit by motivating herbivores to move to their neighboring competitors. Received: 25 March 1999 / Accepted: 8 October 1999     

GEOGRAPHIC PATTERNS OF VARIATION IN OFFSHORE SPOTTED DOLPHINS (STENELLA ATTENUATA) OF THE EASTERN TROPICAL PACIFIC OCEAN

The spotted dolphin (Stenella attenuata), a species commonly associated with the tuna fishery, is widely distributed in the eastern tropical Pacific Ocean (ETP). Geographic patterns in morphological characteristics of this dolphin were evaluated for 36 skull measurements of 574 museum specimens allocated to 5° latitude-longitude blocks. A strong subdivision between northern and southern offshore populations was demonstrated using principal components, canonical variates and cluster (UPGMA and function-point) analyses of block means derived from standardized, sex-adjusted measurements. Classification functions derived from a stepwise multiple discriminant analysis correctly identified more than 87 percent of the specimens as being from either a northern or southern locality. These results support recognition of southern spotted dolphins as a separate management stock. Dolphins from a sample representing a far-western locality show similarities with those from the south. Patterns of geographic variation in morphology were evaluated using Mantel tests and matrix correlations; 17 of the 36 characters showed regional patterning and 18 exhibited local patterning. Morphological trends were also assessed with respect to 14 oceanographic measures that characterize environmental variability in the ETP. In general, clinal variation in morphology mirrors a north-south trend found in several environmental variables. Overall, the most striking associations between morphological and environmental variables involved Solar Insolation (Jan.). Sea surface temperatures also exhibited a strong association with morphological variables; the highest correspondence was found between width of the temporal fossa and sea surface temperature in July. In addition, the thickness of the oxygen minimum layer exhibited a pattern of variation that was statistically associated with 58 percent of the skull characters. The geographic patterns found for cranial morphology and environmental measures are strongly clinal and in general congruent, suggesting that oceanographic conditions in part determine trends in morphological characteristics of spotted dolphins.     

Cytoskeleton-membrane interactions in the cnidocil complex of hydrozoan nematocytes

Summary Each cnidocil complex of the hydrozoans Tubularia larynx and Hydra vulgaris consists of 9 or 7–10 large stereovilli (=stereocilia), respectively, and a modified cilium, the cnidocil. The cnidocils comprise the regular 9 microtubule doublets, up to 30 additional microtubules, as well as a central filament body. Adjacent stereovilli are linked together by intermembrane connectors forming the stereovillar cone. The distal tips of the stereovilli surround the cnidocil in a closed tubular arrangement measuring up to 0.7 m in length. Within this contact region the cnidocil is linked to the stereovillar tube by another set of intermembrane connectors, which seem to hold the cnidocil in a central position within the stereovillar cone. Stereovillar membrane and actin core are linked by 16-nm long cross bridges, which display a periodicity of 16 nm and emerge from the actin core. Within the cnidocils periodically arranged membrane-cytoskeleton bridges are uniformly restricted to the contact region. Here, 24-nm long cross bridges, which are spaced by a regular distance of 20 nm, interconnect the A-tubules of the microtubule doublets and the membrane. The cnidociliary membrane is differentiated into distinct domains as revealed by freeze-fracturing. Within the contact region of the nematocytes of Tubularia larynx, intramembrane particles are arranged in 9 rows of 700 nm length and 50 nm width, separated by particlefree areas. Intramembrane particles are irregularly distributed distal to the contact region. Considering recent physiological results we presume that the latter represent chemoreceptor units, while mechanical stimuli are transmitted via the intermembrane connectors and the microtubule-membrane bridges to mechanosensitive channels within the domain of the cnidociliary membrane in the contact region.     

A scanning electron microscopic investigation of the outer surfaces of Gorgoderina attenuata

The external surface of Gorgoderina attenuata is characterized by ridges that encircle the worm at regular intervals. On the ridges are protuberances that give a beaded appearance to the tegumental surface. Ridges running perpendicular to the encircling bands are apparent close to both suckers. The raised protuberances of the tegumental areas may serve to increase the absorptive surface. A pit-like cavity was observed on the dorsal surface of the oral sucker. In addition numerous raised structures which appear to be of two types were present on the surface of the oral sucker. One type resembles a depression with a small knob barely visible in the center and the second type consists of button-like patches of tissue. From their structure and location these are tentatively identified as sensory organs.     

广西润楠属一新种－狭基润楠

报道了润楠属一新种,狭基润楠Machilus attenuataF.N.Wei&S.C.Tang。本种与形态上相似的定安润楠M.dinganensis S.Lee&F.N.Wei的区别在于叶先端尾状渐尖或长渐尖,基部渐狭并略下延,幼叶上面无毛,下面疏被短柔毛;与西畴润楠M.sichourensisH.W.Li的区别在于叶较小,薄革质,倒披针形或椭圆形,长8~13 cm,宽2.2~4 cm,基部渐狭,侧脉在上面明显或略明显,下面明显。     

灰麝鼩的种群数量变动

汇集了1989～1996年云南保山市坝区的调查资料和其它地区的文献资料,对灰麝鼩种群数量变动进行了分析,灰麝鼩集中分布于海拔1650m的保山市坝区,具有较强的区域性和海拔高度的局限性特征.种群相对数量高峰分别出现在6月和8～12月.灰麝鼩的年变动在保山坝区居民区鼠型小兽群落中组成相对稳定.     

Herbivore-induced plant vaccination. Part I. The orchestration of plant defenses in nature and their fitness consequences in the wild tobacco Nicotiana attenuata

A plant's responses to attack from particular pathogens and herbivores may result in resistance to subsequent attack from the same species, but may also affect different species. Such a cross-resistance, called immunization or vaccination, can benefit the plant, if the fitness consequences of attack from the initial attacker are less than those from subsequent attackers. Here, we report an example of naturally occurring vaccination of the native tobacco plant, Nicotiana attenuata, against Manduca hornworms by prior attack from the mirid bug, Tupiocoris notatus (Dicyphus minimus), which results from the elicitation of two categories of induced plant responses. First, attack from both herbivore species causes the plants in nature to release predator-attracting volatile organic compounds (VOCs), and the attracted generalist predator, Geocoris pallens, preferentially attacks the less mobile hornworm larvae. Second, attack from both mirids and hornworms increases the accumulation of secondary metabolites and proteinase inhibitors (PIs) in the leaf tissue, which is correlated with the slow growth of Manduca larvae. Mirid damage does not significantly reduce the fitness of the plant in nature, whereas attack from the hornworm reduces lifetime seed production. Consequently, plants that are attacked by mirids realize a significant fitness advantage in environments with both herbivores. The combination of growth-slowing direct defenses and predator-attracting indirect defenses results in greater hornworm mortality on mirid-attacked plants and provides the mechanism of the vaccination phenomenon.     

DIFFERENCES BETWEEN FISHERY-DEPENDENT AND FISHERY-INDEPENDENT ESTIMATES OF SINGLE-AND MIXED-SPECIES DOLPHIN SCHOOLS: IMPLICATIONS FOR SINGLE-SPECIES STOCK ASSESSMENTS

Since 1979, fishery-independent data and fishery-dependent data have been used to estimate trends in the abundance of the northeastern stock of spotted dolphins in the eastern tropical Pacific. Data collected aboard tuna vessels have been used to estimate trends in relative abundance, while data collected from fishery-independent research vessels have been used to estimate trends in actual abundance. One of the largest discrepancies between the two data sources is that tuna vessels tend to report dolphin schools that are 400%—500% larger than schools observed by research vessels. After comparing research vessel and tuna vessel observations overlapping in space and time, it appears that either measurement error or selective reporting of large schools is the most likely explanation for the disparity. Comparing single-species schools against portions of mixed- species schools introduced bias. Revised fishery-dependent abundance estimates were generated using only single-species dolphin schools, resulting in estimates that appear more similar to recent fishery-independent abundance estimates.
correctable in fishery-dependent data, suggesting that data on mixed-species schools should not be considered if fishery-dependent data are to be combined with fishery-independent data for stock assessment purposes.     

Effect of a saprophytic fungus on the growth and the lead uptake, translocation and immobilization in Dodonaea viscosa

Phytoremediation is a feasible alternative to remediate soils polluted with toxic elements, which can be enhanced by manipulating plant-microbe interactions. Regarding this, free-living saprophytic fungi that interact beneficially with roots have been scarcely studied. Thus, the aim of this study was to assess the effect of a saprophytic fungus, Lewia sp., on the plant growth and the ability of Dodonaea viscosa to phytoaccumulate or phytostabilize soluble and insoluble sources of lead in a solid support. The growth of D. viscosa was influenced by both Pb and Lewia sp. While seedlings exposed to Pb showed a decrease in biomass production, in seedlings grown without Pb the biomass was stimulated by Lewia sp. The fungus strongly stimulated the weight-to-length ratio in roots. Regardless of the treatment, D. viscosa accumulated 4.4-6.5 times more Pb in roots than in shoots, conducting to low translocation factors (< 0.2). The presence of Lewia sp. significantly improved Pb accumulation, achieving high bioconcentration factors (> 22), which was attributed to an increased bioavailability and uptake of Pb due to the fungus. This study demonstrated that Lewia sp. could improve Pb-phytostabilization by D. viscosa in soils polluted with soluble and insoluble forms of Pb.