Timing of exposure to ozone affects reproductive sensitivity and compensatory ability in Brassica campestris

It is well established that exposure to ozone (O₃) may impair vegetative growth and reproductive development in plants, although the consequences for yield depend on the effectiveness of the compensatory processes induced. This study examined the effects of exposing the terminal inflorescence of Brassica campestris L. to 100 ppb O₃ for 6 h d⁻¹ on four consecutive days during early flowering while the vegetative organs received charcoal-filtered air. The ordered predictability of development in B. campestris is ideal for studies of the impact of abiotic stress factors such as O₃ on reproductive development and seed production. Effects on reproductive development and seed yield characteristics were determined for floral sites exposed at different developmental stages. Flower and pod numbers on the terminal raceme were unaffected by exposure, but effects on pod length varied depending on the developmental stage of floral sites during exposure. Increased ovule abortion and precocious seed germination in the pods of O₃-treated plants reduced mature seed number pod⁻¹. Although the individual weight of mature seeds was slightly greater in O₃-treated plants, seed yield pod⁻¹ and seed yield plant⁻¹ were reduced due to the lower seed number pod⁻¹. Seed from O₃-treated plants germinated more rapidly than control seed irrespective of the stage of floral development during exposure. The results show that exposure to realistic O₃ episodes during the early stages of flowering significantly reduced seed yield without impairing the vigour of the seeds produced. The physiological origins of these effects are discussed.     

不同结荚习性夏大豆种质的农艺表现及其与产量的相关分析

以533份不同结荚习性大豆种质为试验材料,研究了不同结荚习性大豆种质在黄淮夏播生态区的农艺性状表现,并对主要农艺性状和产量的相关性进行了分析。结果表明:大豆种质的营养期、株高、有效分枝、单株荚数、倒伏性、株型等性状的平均值随无限-亚有限-有限结荚习性递减,生殖期、单株粒重、百粒重、小区产量等性状平均值的变化趋势则相反。相关分析表明,无限结荚习性种质的产量与株高、单株粒重呈极显著正相关,偏相关系数分别为0.602**、0.566**,与有效分枝、倒伏性呈显著负相关,偏相关系数分别为-0.384*、-0.451*。亚有限结荚习性种质的产量与生殖期、单株粒重呈显著、极显著正相关,偏相关系数分别为0.156*、0.536*,与有效分枝呈极显著负相关,偏相关系数为-0.323**。有限结荚习性种质的产量与单株粒重、株高呈极显著正相关,偏相关系数分别为0.433**、0.262**,与株型、单株荚数呈显著、极显著负相关,偏相关系数分别为-0.149*、-0.198**。结合不同结荚习性品种的生长特点,本研究认为,无限结荚习性品种株高较高且株高与产量呈极显著正相关,适合干旱地区种植;亚有限结荚习性品种生殖期与产量呈正相关,生殖期内生长旺盛需要较多的养分供应;有限结荚习性品种的营养生长持续时间短,株高较矮,吸收光能有限,实现高产主要依赖各性状器官间的平衡。生产中,有限结荚习性品种的营养生长期间既需要充足的肥水促其生长,又要防止旺长。     

SOYBEAN GROWTH RESPONSES TO ENHANCED LEVELS OF ULTRAVIOLET-B RADIATION UNDER GREENHOUSE CONDITIONS

Soybean (Glycine max [L.] Merr. cv. Essex) was grown in an unshaded greenhouse under three levels of biologically effective ultraviolet-B (UV-B_BE) radiation (effective daily dose: 0, 11.5 and 13.6 kJ m^–2) for 91 days. Plants were harvested at regular intervals beginning 10 days after germination until reproductive maturity. Mathematical growth analysis revealed that the effects of UV-B radiation varied with plant growth stage. The transition period between vegetative and reproductive growth was the most sensitive to UV-B radiation. Intermediate levels of UV-B had deleterious effects on plant height, leaf area, and total plant dry weight at late vegetative and reproductive stages of development. Specific leaf weight increased during vegetative growth but was unaffected by UV-B during reproductive growth stages. Relative growth, net assimilation, and stem elongation rates were decreased by UV-B radiation during vegetative and early reproductive growth stages. Variation in plant responses may be due in part to changes in microclimate within the plant canopy or to differences in repair or protection mechanisms at differing developmental stages.     

Short-term UV-B radiation and ozone exposure effects on aromatic secondary metabolite accumulation and shoot growth of flavonoid-deficient Arabidopsis mutants

The presence of UV-absorptive substances in the epidermal cells of leaves is thought to protect mesophyll tissues from the harmful effects of UV-B radiation. We examined the influence of short-term UV-B exposures on UV-absorptive (330 nm) sinapates and flavonols, and on shoot growth of the Arabidopsis wild type ecotype Landsberg erecta and two mutants. 114 deficient in chalcone synthase, and 115 , deficient in chalcone/flavonone isomerase. Sequential ozone exposures were used to determine the effects of oxidative stress The levels of sinapates and flavonols on a leaf fresh weight basis increased substantially in the wild type and sinapates increased in the 114 mutant in vegetative vegetative/reproductive transitional and reproductive stage plants in response to short-term (48h) UV-B radiation. When UV-B was discontinued the levels generally decreased lo pre-exposure levels after 48 h in vegetative/reproductive but not in reproductive plants. Exposure to ozone before or alter UV-B treatment did not consistently affect the levels of these UV-absorptive compounds. Dry matter accumulation was less affected by UV-B at the vegetative and reproductive stages than at the vegetative/reproductive stage. At the vegetative/reproductive stage, shoot growth of all 3 genotypes was retarded by UV-B. Growth was not retarded by short-term ozone exposure alone but when exposure to ozone followed UV-B exposure, growth was reduced in all genotypes. Leaf cupping appeared on 115 plants exposed to UV-B.     

Ineffective utilization of nitrate by soybean during pod fill

The relative effectiveness of nitrate, allantoin, or nitrate plus allantoin as sources of nitrogen for the indeterminate soybean plant [ Glycine max (L.) Merr cv. Harper] was studied throughout vegetative and reproductive growth. All plants were provided with 3.0 m M nitrogen and were grown hydroponically in growth chambers. During vegetative and early reproductive growth, plants given nitrate or nitrate plus allantoin grew faster than plants provided allantoin only. However, during pod fill, plants provided with allantoin or allantoin plus nitrate gained weight more rapidly than plants receiving just nitrate. More importantly, at maturity plants that had been provided with allantoin or allantoin plus nitrate during pod fill were 30% heavier in total dry weight, 50% higher in nitrogen content, and 50% higher in seed yield than plants that had received just nitrate. At full bloom, all plants were inoculated with the same culture of Bradyrhizobium japonicum , and twice each week throughout pod fill each plant was assayed for nitrogen fixation (acetylene reduction). Correlation coefficients obtained by linear regression analysis show a strong positive correlation between the measured rate of nitrogen fixation and maximum plant fresh weight (r = 0.83), total plant nitrogen (r = 0.81), or seed yield (r = 0.76). The fact that nitrogen fixation during pod fill stimulates plant growth and seed yield, coupled with the facts that nitrate blocks nodulation and is not used efficiently during pod fill by the soybean plant, may explain why seed yield of field-grown soybeans usually does not respond to added fertilizer nitrogen. Thus, it is suggested that enhanced nitrogen fixation may be the key factor in improving soybean seed yield.     

The role of reproductive plant traits and biotic interactions in the dynamics of semi-arid plant communities

The dynamics of semi-arid plant communities are determined by the interplay between competition and facilitation among plants. The sign and strength of these biotic interactions depend on plant traits. However, the relationships between plant traits and biotic interactions, and the consequences for plant communities are still poorly understood. Our objective here was to investigate, with a modelling approach, the role of plant reproductive traits on biotic interactions, and the consequences for processes such as plant succession and invasion. The dynamics of two plant types were modelled with a spatially-explicit integrodifferential model: (1) a plant with seed dispersal (colonizer of bare soil) and (2) a plant with local vegetative propagation (local competitor). Both plant types were involved in facilitation due to a local positive feedback between vegetation biomass and soil water availability, which promoted establishment and growth. Plants in the system also competed for limited water. The efficiency in water acquisition (dependent on reproductive and growth plant traits) determined which plant type dominated the community at the steady state. Facilitative interactions between plant types also played an important role in the community dynamics, promoting establishment in the driest conditions and recovery from low biomass. Plants with vegetative propagation took advantage of the ability of seed dispersers to establish on bare soil from a low initial biomass. Seed dispersers were good invaders, maintained high biomass at intermediate and high rainfall and showed a high ability in taking profit from the positive feedback originated by plants with vegetative propagation under the driest conditions. However, seed dispersers lost competitiveness with an increasing investment in fecundity. All together, our results showed that reproductive plant traits can affect the balance between facilitative and competitive interactions. Understanding this effect of plant traits on biotic interactions provides insights in processes such as plant succession and shrub encroachment.     

Relationship between the photosynthetic activity and the performance of Cakile maritima after long-term salt treatment

Cakile maritima is a halophyte with potential for ecological, economical and medicinal uses. We address here the impact of salinity on its growth, photosynthesis and seed quality. Whole plant growth rate and shoot development were stimulated at moderate salinity (100–200 m M NaCl) and inhibited at higher salt concentrations. Although diminished in the presence of salt, potassium and calcium uptake per unit of root biomass was maintained at relatively high value, while nutrient-use efficiency (NUE) was improved in salt-treated plants. Chl and carotenoid concentrations decreased at extreme salinities, but anthocyanin concentration continuously grew with salinity. Net photosynthetic rate (A), stomatal conductance, maximum quantum efficiency of PSII and quantum yield were stimulated in the 100–200 m M NaCl range. Higher salinity adversely affected gas exchange and changed PSII functional characteristics, resulting in a reduction of A per leaf area unit. This phenomenon was associated with increased non-photochemical quenching. Harvest index, silique number and seeds per fruit valve were maximal at 100 m M NaCl. Despite the decreasing salt accumulation gradient from the vegetative to the reproductive organs, high salinities were detrimental for the seed viability and increased the proportion of empty siliques. Overall, the salt-induced changes in the plant photosynthetic activity resulted into analogous responses at the vegetative and reproductive stages. The enhancement of NUE, the absence of pigment degradation, the reduction of water loss and the concomitant PSII protection from photodamage through thermal dissipation of excess excitation significantly accounted for Cakile survival capacity at high salinity.     

The influence of pre-floral growth and development on the pathway of floral development, dry matter distribution and seed yield in oilseed rape (Brassica napus L.)

Patterns of floral development, dry matter distribution and seed yield were examined in winter oilseed rape plants subjected to different pre-floral growth environments. The duration of pre-floral growth and plant size at flower initiation, measured in terms of total mainstem leaf number, were manipulated by varying the temperature between seedling emergence and flower initiation. Exposure of seedlings to low temperatures from cotyledon expansion onwards markedly reduced the duration of pre-floral growth and the number of leaves on the mainstem. The subsequent development pattern of plants was largely dependent on the date of flower initiation and therefore vernalisation requirement. Indeed, the period of growth from flower initiation to maturity, considered on the basis of thermal time, was directly related to the duration of pre-floral growth and mainstem leaf number. The thermal durations of the bud development phase and flowering period in plants exposed to different pre-floral cold treatments but with a common date of flower initiation were similarly linked to these two parameters. Plants exposed to prolonged periods of low temperature treatment from cotyledon expansion onwards initiated fewer mainstem leaves during a relatively short pre-floral growth phase and their yield potential was limited by a reduction in branch and flower numbers. Plants maintained at higher temperatures produced more mainstem leaves during an extended period of pre-floral growth and supported a greater number of branches and flowers. However, this additional yield potential was not realised due to a reduction in seed numbers and mean seed weight. It appeared that seed yield of these plants was limited by increased competition between an excessive number of lower branches and flowers, a problem apparently created by excessive pre-floral growth. Minimal competition for available assimilates between the limited number of branches of plants with a shorter pre-floral growth phase and fewer mainstem leaves, resulted in lower levels of pod abortion, greater seed production and ultimately increased seed yields.     

From over to undercompensation: Variable responses to herbivory during ontogeny of a Neotropical monocarpic plant

Empirical and theoretical work has suggested that plants can change their compensatory responses to herbivory as they develop. However, such a relationship is likely to be more complex than previously thought since the amount and type of damage a plant receives can also change as the plant develops. Here, we determined the survival, growth, and reproductive output of plants (Actinocephalus polyanthus) from different ontogenetic stages that received variable levels of natural or simulated herbivore damage. Juvenile plants and non‐reproductive adults in which leaves were damaged showed full vegetative compensation, whereas pre‐reproductive plants were not able to replace the lost leaves. However, these same pre‐reproductive plants produced more inflorescences and thus more seeds and seedlings than control plants. In contrast, damage to vegetative and/or reproductive structures during the reproductive phase resulted in a negative effect on seed and seedling production. Herbivory effects on plant survival, growth, and reproduction during the vegetative and pre‐reproductive phases were independent of the amount of damage. However, during reproduction, the magnitude of these effects was strongly influenced by the amount of damage and the reproductive stage of the plant at the time of the damage. In short, our results demonstrate that the survival, growth, and reproductive responses to herbivory of A. polyanthus can be dependent on the timing and/or intensity of damage. The reproductive response of A. polyanthus to our simulated herbivory treatments during the pre‐reproductive phase represents an example of overcompensation. Furthermore, it indicates that vegetative regrowth is not necessarily a driving factor for tolerance.     

农田冬小麦生长和产量对臭氧动态暴露的响应

 评估臭氧(O₃)污染对农田冬小麦生长和产量的影响是污染生态学和生理生态学研究的重要内容之一。该研究运用开顶式气室(OTC),对冬小麦‘ 嘉403’(Triticum aestivum cv. Jia 403)进行了O₃动态暴露的田间原位试验。实验设置过滤空气组(CF)、自然大气组(NF)和两个不同浓度的 O₃动态暴露组(DO100和DO150)。结果表明：1) O₃浓度增加,一方面可以改变灌浆期冬小麦叶片气体交换参数的日变化规律;另一方面引起表观 光量子产额、光饱和点和光补偿点等光响应参数的显著降低。这表明灌浆期叶片光合能力的下降是气孔因素和非气孔因素共同作用的结果。2) O₃暴露可以改变小麦形态特征,如植株变矮、叶片衰老加速、 叶面积变小,并最终导致产量大幅下降。     

Growth and gas exchange response to water shortage of a maize crop on different soil types

The effect of water shortage on growth and gas exchange of maize grown on sandy soil (SS) and clay soil was studied. The lower soil water content in the SS during vegetative growth stages did not affect plant height, above-ground biomass, and leaf area index (LAI). LAI reduction was observed on the SS during the reproductive stage due to early leaf senescence. Canopy and leaf gas exchanges, measured by eddy correlation technique and by a portable photosynthetic system, respectively, were affected by water stress and a greater reduction in net photosynthetic rate (A _N) and stomatal conductance (g _s) was observed on SS. Chlorophyll and carotenoids content was not affected by water shortage in either condition. Results support two main conclusions: (1) leaf photosynthetic capacity was unaffected by water stress, and (2) maize effectively endured water shortage during the vegetative growth stage.     

Multiple switches between vegetative and reproductive growth in annual plants

A model of growth and reproduction in annual plants was developed by Cohen (1971, J. Theor. Biol.33, 299–307) to determine the allocation strategy which maximizes seed yield. The model divides the plant into vegetative and reproductive parts and predicts that yield is maximized by a strategy consisting of a switch from purely vegetative to strictly reproductive growth. We generalize Cohen's model to include vegetative and reproductive loss terms. Both growth and loss rates are allowed to vary with time. Using optimal control theory we find that seed yield is maximized by a strategy consisting of multiple switches between vegetative and reproductive growth, for certain ranges of the model parameters. In natural systems a predictable vegetative loss burst may be necessary to promote multiple switches.     

Effect of enhanced UV-B radiation on pollen quantity,quality, and seed yield in Brassica rapa (Brassicaceae)

We conducted three experiments to examine the influence of ultraviolet-B radiation (UV-B; 280–320 nm) exposure on reproduction in Brassica rapa (Brassicaceae). Plants were grown in a greenhouse under three biologically effective UV-B levels that simulated either an ambient stratospheric ozone level (control), 16% (“low enhanced”), or 32% (“high enhanced”) ozone depletion levels at Morgantown, WV, USA in mid-March. In the first experiment, we examined whether UV-B level during plant growth influenced in vivo pollen production and viability, and flower production. Pollen production and viability per flower were reduced by ≈50% under both enhanced UV-B levels relative to ambient controls. While plants under high-enhanced UV-B produced over 40% more flowers than plants under the two lower UV-B treatments, whole-plant production of viable pollen was reduced under high-enhanced UV-B to 17% of that of ambient controls. Whole-plant production of viable pollen was reduced under low-enhanced UV-B to 34% of ambient controls. In the second experiment, we collected pollen from plants under the three UV-B levels and examined whether source-plant UV-B exposure influenced in vitro pollen germination and viability. Pollen from plants under both enhanced-UV-B treatments had initially lower germination and viability than pollen from the ambient level. After in vitro exposure to the high-enhanced UV-B levels for 6 h, viability of the pollen from plants grown under ambient UV-B was reduced from 65 to 18%. In contrast, viability of the pollen from plants grown under both enhanced UV-B treatments was reduced to a much lesser extent: only from ≈43 to 22%. Thus, ambient source-plant pollen was more sensitive to enhanced UV-B exposure. In the third experiment, we used pollen collected from source plants under the three UV-B levels to fertilize plants growing under ambient-UV-B levels, and assessed subsequent seed production and germination. Seed abortion rates were higher in plants pollinated with pollen from the enhanced UV-B treatments, than from ambient UV-B. Despite this, seed yield (number and mass) per plant was similar, regardless of the UV-B exposure of their pollen source. Our findings demonstrate that enhanced UV-B levels associated with springtime ozone depletion events have the capacity to substantially reduce viable pollen production, and could ultimately reduce reproductive success of B. rapa.     

COMPONENTS OF REPRODUCTIVE EFFORT AND YIELD IN GOLDENRODS

Four components of reproductive yield (the weight of reproductive tissue) were examined in relation to their effect on reproductive effort and their relative contributions to reproductive yield in five species of goldenrods (Solidago, Compositae). The yield components were number of flowing stems per plant, number of flowering branches per stem, number of flowering heads per branch, and number of seeds per seed head. Individuals within populations increase their reproductive effort by increasing their reproductive weight, not by decreasing their vegetative weight. Each species shows a different pattern of positive correlations of yield components with reproductive yield and reproductive effort, indicating that each species has its own mechanisms for regulating reproduction. The yield components were not significantly intercorrelated.     

A GmAOX2b antisense gene compromises vegetative growth and seed production in soybean

The alternative oxidase mediates the cyanide-resistant respiratory pathway in plant mitochondria. In non-thermogenic plants, the role of alternative oxidase in plant growth and development is not well understood. Soybean (Glycine max) lines carrying a GmAOX2b antisense gene had compromised vegetative growth and reproductive performance under typical glasshouse growth conditions. The reduction in vegetative growth was demonstrated by reduction in shoot height, the number of leaves per plant and the green leaf area. Antisense plants also had decreased pod formation and seed to pod ratios, which together led to a reduction in the number and total mass of seed produced. The negative effects of the antisense gene on pod set, seed set, ovule availability and total seed mass were primarily confined to the branches, rather than the main stem. The preferential effect of alternative oxidase suppression in the branches is discussed in relation to the reproductive potential of soybean under stress. Taken together, these results demonstrate that alternative oxidase provides the benefit of sustaining plant vegetative growth and reproductive capacity in soybean.     

EFFECTS OF FIRE IN TALLGRASS PRAIRIE ON GROWTH AND REPRODUCTION OF PRAIRIE CONEFLOWER (RATIBIDA COLUMNIFERA: ASTERACEAE)

Effects of fire on growth and reproduction of the perennial forb Ratibida columnifera were studied on the Konza Prairie Research Natural Area in northeastern Kansas, USA. Populations were sampled in seven different tallgrass prairie watersheds that varied in fire frequency and in the number of years elapsed since the last fire. Plants from sites not burned for many years were 2.6 times larger and produced 50% more stems than plants from recently burned sites. Number of seeds per plant was also higher in long-unburned sites due to greater numbers of flower heads per plant and greater numbers of achenes produced per head. Reproductive effort (ratio of inflorescence biomass to total vegetative biomass) was 33% lower in annually burned prairie than in any of the other sites. Significant differences in the relationships of inflorescence biomass to vegetative plant biomass in burned vs. unburned sites indicated that burning causes direct changes in plant reproductive effort independent from its effects on plant size. There was no clear relationship between patterns of seed production among sites and patterns of R. columnifera abundance. Ratibida columnifera responses to fire are most likely a result of changes in the relative competitive abilities of forbs and the dominant perennial grasses due to post-fire changes in abiotic conditions rather than a result of direct effects of fire on the fate of buds and subsequent vegetative and floral development.     

Reproductive Growth and Dry Matter Production of Glycine max (L.) Merr. in Response to Oxygen Concentration

Reproductive as well as vegetative parameters of mature soybean (Glycine max [L.] Merr. cv. Wye) plants grown in chambers in which the aerial portion was exposed to altered pO₂ during all or part of the growth cycle were measured. Oxygen concentration was found to be a key factor controlling all phases of reproductive development. Exposure to 5% O₂ from early seedling stage to senescence increased leaf, stem, and root dry weights and reduced seed yields when compared to 21% O₂; exposure to low O₂ during the vegetative growth stage from early seedling to mid-flowering arrested pod but not seed development; exposure from mid-flowering to mid-pod filling almost completely arrested seed but not pod development; exposure from mid-pod filling to senescence arrested seed development at the mid-filling stage.     

Lolium multiflorum density responses under ozone and herbicide stress

Adaptations to overcrowding of individual plants result in density dependant control of growth and development. There is little information on how anthropogenic stresses modify these responses. We investigated whether combinations of diclofop‐methyl herbicide and tropospheric ozone alter the pattern of expected growth compensation with density changes resulting from intraspecific competition in Lolium multiforum Lam (Poacea) plants. Individual plant vegetative parameters and total seed production were assessed for plants growing under various densities and different herbicide rates and ozone treatments. The stressors differently changed the frequency distribution for average individual plant weight resulting from increasing densities. Only herbicide affected seedling mortality. Plants were able to compensate during grain filling maintaining similar seed production – density relationships in all treatments. Our findings contribute to the understanding of the impact of stress factors on the demographic changes in plant populations. Important ecological implications arise: (i) contrasting responses to ozone and herbicide, alone and in combination of individual plants resulted in different biomass – density relationships; (ii) stress effects on plant populations could not be predicted from individual responses; and (iii) changes in competitive outcome by single or combined stress factors may alter the expected genotype frequency in a crowded population with few dominant individuals.     

The response of an autumn-sown determinate faba bean to different plant distributions

The response of an autumn-sown determinate selection, 858 , to different plant distribution patterns was examined in field trials during the two seasons 1985/86 and 1986/87 at the University of Nottingham. Plants were sown at two densities (20 and 40 plants per m2) and at three different row widths (11.9, 23.8 and 47.6 cm) in the autumn of each season. Plant numbers and combined yields were greater in 1986/87 than in 1985/86. Grain yield was unaffected by differences in inter-row spacing and this was ascribed to the early attainment of complete ground cover even when plants were grown in widely spaced rows. Yield was most strongly correlated with seed numbers per m². An examination of yield on a per plant basis suggested that narrow inter-row spacings were more productive at low plant densities, while yield per plant was greater at higher densities if the crop was grown on wider inter-row spacings. It was concluded that autumn-sown determinate forms of faba bean were able to compensate for large changes in plant distribution and consequently yield was unaffected.     

Opium poppy capsule growth and alkaloid production is constrained by shade during early floral development

Morphinan alkaloids accumulate in the capsules of Papaver somniferum L. (opium poppy) most likely as defence against herbivory. Thus, capsule size is an important component of alkaloid yield. Shade during early cell division-dominated growth of reproductive structures generally reduces final fruit size more than shade during later cell expansion-dominated growth. The current study aimed to determine whether this response is found in opium poppy and the subsequent impact on alkaloid yield, composition and seed production. First the timing of key reproductive developmental events was resolved relative to macromorphological traits. Plants were then shaded during either (a) floral initiation, (b) early floral development or (c) capsule expansion before being harvested at maturation. Shade during floral initiation dramatically reduced final capsule size, alkaloid yield and seed number, and increased the concentration of precursor compounds relative to morphine, despite plants later returning to full sun. Shade during later capsule growth enhanced capsule size and alkaloid yield but had little effect on alkaloid composition or seed number. Thus, early developmental processes, including morphine biosynthesis, appear to have a relatively greater demand for carbohydrates compared with later processes. Crop management practices and environmental factors that limit carbohydrate availability during early development are thus predicted to have significant negative impacts on alkaloid production and reproductive success.