Precipitation-dependent source–sink dynamics in a spatially-structured population of an outbreaking caterpillar

Context

Patch-based population models predominately focus on factors that affect regional processes namely, patch size and connectivity, as the primary drivers explaining patch occupancy. This trend persists despite the recognition that patch quality can strongly influence population demography at the local scale. The quality of patches is often temporally variable and influenced by abiotic conditions. However, few studies have explicitly investigated how climatic variables influence the spatial and temporal dynamics of spatially-structured populations either directly or indirectly through changes in patch quality.

Objectives

Using a 10-year census of a spatially-structured population of an outbreaking caterpillar, we determined the relative importance of patch quality (determined demographically), connectivity, precipitation, and their interactive effects on patch abundance, occupancy, colonization, and extinction.

Methods

We generated a series of statistical models and performed comparisons using Akaike’s information criterion. We subsequently used likelihood ratio tests to determine the influence of each parameter on model fit.

Results

Patch quality and precipitation were the strongest predictors of the observed dynamics. We found that the dynamics of the spatially-structured population of Arctia virginalis were strongly influenced by precipitation: all patches had a higher probability of occupancy, contained higher abundances of caterpillars, and experienced fewer extinctions following wet winters compared to years following droughts.

Conclusion

These findings suggest that precipitation may act to influence the strength of heterogeneity of patch quality. This work demonstrates that patch-based models that do not include local and climatic factors may produce poor predictions under future climatic regimes.

     

Effect of nitrogen and calcium stock plant nutrition on Petunia × hybrida leaf and anther explant growth in vitro

Petunia × hybrida Hort. Vilm.-Andr. ‘Coral Sea’ stock plants were grown with a modified Hoagland's solution to determine the effect of stock plant N and Ca nutrition on subsequent leaf explant and anther growth in vitro. In separate experiments, N was 0, 75, 150 or applied at 0, 100, 200 or 400 mg 1⁻¹ as nitrate-N, and Ca was applied at 300 mg 1⁻¹. Leaf explants from plants grown with 100 mg 1⁻¹ N produced the greatest amount of callus, while those from plants grown with 200 and 400 mg 1⁻¹ N produced the highest number of shoots. Nitrogen treatments increased the anther fresh weight and the number of shoots per culture compared to anthers derived from stock plants that did not receive N.The fresh weight of shoots produced by leaf explants decreased with increasing Ca concentrations applied to stock plants. Anthers derived from stock plants treated with 0 mg 1⁻¹ Ca produced the highest number of shoots per culture.Anatomically, both filament and anther wall tissue produced callus, but no callus production from microspores was observed. Organs produced by anthers were diploid.     

The effect of boron upon leaf development and growth of the apple cultivar ‘Cox’s Orange Pippin’

One-year-old ‘Cox’s Orange Pippin’ trees planted in pots in sand culture in the spring of 1970 were (during the growing seasons of 1970–72) watered with nutrient solutions containing 0, 0.12, 0.23, 1.2, 3.5 or 7.0 mg boron per litre.

With increasing boron supply, the boron concentration of the leaves increased, the growth of current year’s shoots and to some extent of other plant parts increased, and summer leaf-fall and the occurrence of necrotic leaf spots (‘Cox spots’) decreased. The percentage of healthy leaves was greatest when boron in leaf dry matter was about 30 ppm.

In 1972 the trees with highest boron supply suffered from incipient boron toxicity and increased incidence of ‘Cox disease’ flowering was delayed, and the percentage of dead buds was increased. In the latter part of the summer the leaves curled upwards, turned purple and became brittle.     

Effect of part-tree flower thinning on fruiting,vegetative growth and leaf photosynthesis in ‘Cox’s Orange Pippin’ apple

Flower clusters were removed at full bloom from ten year old ‘Cox’s Orange Pippin’ trees on M.9 rootstocks, over the whole tree, on alternate branches or on whole sides. Mean fruit weight per tree at harvest was linearly dependent on leaf area per fruit and on light interception per fruit, both relationships accounting for over 90% of the variance. These relationships did not differ between treatments, implying either a mobile pool of carbohydrate or photosynthetic adjustment within the tree to crop load. Measurements of leaf photosynthesis in July and September showed no statistically significant differences in photosynthetic rate of spur or extension shoot leaves on bearing or non-bearing branches. Although the treatments caused no overall effects on shoot growth or leaf area per tree, sides of trees without fruit had greater leaf area and shoot growth than did sides bearing fruit. Fruit mineral composition and percentage dry weight were not affected by treatment except where the treatments significantly altered fruit size. In the following spring, although the treatments did not affect the total number of flower buds produced, branches that were deflowered in the previous spring carried significantly more flower buds than did branches which had cropped.     

Culture method and photosynthetic photon flux affect photosynthesis,growth and survival of Limonium ‘Misty Blue’ in vitro

Microcuttings (shoots each with two leaves) of Limonium ‘Misty Blue’ were cultivated in vitro for 28 days under photoautotrophic (sucrose-free culture medium; CO₂ and photosynthetic photon flux (PPF) enriched conditions), photomixotrophic (medium with 30 g l⁻¹ sucrose; CO₂ and PPF enriched conditions) and heterotrophic (medium with 30 g l⁻¹ sucrose; CO₂ non-enriched conditions) methods. Several growth variables were measured during and at the end of cultivation: shoot fresh and dry weight, percentage of shoot dry matter, root fresh weight, number of leaves, leaf area, chlorophyll and sugar content of leaves, stomatal density and size, net photosynthetic rate (NPR) and percent survival of plantlets ex vitro. Plantlets grown in photoautotrophic and photomixotrophic methods had more leaves, high chlorophyll and sugar contents, high NPR, and showed high percent survival. However, these plantlets possessed less number of stomata per square millimeter. In contrast, the plantlets grown by the heterotrophic method showed decreased values of these growth variables except for the number of stomata per square millimeter. These results indicate that CO₂ enrichment for plantlets in vitro at a relatively high PPF would promote photosynthesis and hence growth of chlorophyllous explants/plantlets in vitro. The resulting plantlets were acclimatized better and sooner on ex vitro transplantation.     

Divergent functions of SNAC4–9 and possible mechanisms for tomato adaptation to abiotic stresses

Plants are exposed to all types of abiotic stresses during the process of growth and development, which could adversely affect the productivity and postharvest storage quality of plants. In the current study, the expression of SNAC4–9 and the changes of physiological parameters were analyzed in tomato seedlings under various abiotic conditions and hormone treatments. The results demonstrate that all six genes were induced by these stresses at differential induction levels and that SNAC4–9 gene expression profiles were likely to be related to the ABA, SA, and MeJA signaling pathways. In addition, all of the stress- and hormone-treated seedlings exhibited significant increases in their proline content and antioxidant enzyme activities. MDA was significantly increased in seedlings exposed to stress and decreased in hormone-treated seedlings. These data collectively suggest that SNAC4–9 might function through ABA, SA, and JA signaling pathways and the regulation of proline and antioxidant systems. This study combines molecular biology and physiology to provide valuable information for further exploring the functional roles of NAC genes in response to environmental stresses and indicates that these genes may exhibit potential for enhancing stress tolerance of transgenic tomato plants and further improving the postharvest storage quality of tomato plants.     

Influence of rootstock on vegetative growth,fruit yield and quality in ‘Cuore di Bue’, an heirloom tomato

Summary

Heirloom tomatoes (Solanum lycopersicum L.) such as ‘Cuore di Bue’ are highly appreciated by consumers for their outstanding quality and flavour. Nowadays, they are often grafted onto vigorous rootstocks in order to overcome several soil-borne diseases. The present study was conducted in 2007 and 2008 in Southern Italy. Plants of ‘Cuore di Bue’, an heirloom ‘oxheart’ tomato, were grown in a greenhouse either as non-grafted plants, or grafted onto two inter-specific (S. lycopersicum S. habrochaites) rootstocks (‘Beaufort F₁’ and ‘Maxifort F₁’) in order to evaluate their effects on vegetative growth, marketable yield, fruit quality, and the sensory properties of ‘Cuore di Bue’ tomato fruit. Growth analysis revealed that ‘Maxifort F₁’ enhanced plant growth, particularly in terms of leaf area and leaf fresh weight. Grafted plants had higher leaf area ratios (by 13%) and higher leaf dry weight fractions (by 18%) compared to non-grafted ‘Cuore di Bue’ plants. Marketable yields increased by 20 – 25% in grafted plants in both years. However, total soluble solids (TSS) contents, titratable acidity (TA), and TSS/TA ratios were not significantly affected by grafting.Vitamin C contents decreased by 14 – 20% in both years in the fruit of plants grafted onto either rootstock. The sensory profiles of fruit were not modified by grafting, although taste panelists expressed a higher preference for purchasing fruit from plants grafted onto ‘Maxifort F₁’. These results confirm that, when using the appropriate rootstock/scion combination, grafting can improve plant growth and the marketable yield of heirloom tomato fruit without reducing the sensory quality of the fruit or its biochemical parameters. However, grafting can reduce vitamin C contents and, thus, nutritional quality.     

Effect of tebuconazole-tolerant and plant growth promoting Rhizobium isolate MRP1 on pea–Rhizobium symbiosis

Seed dressing with fungicides adversely affects the structure and function of beneficial soil microbial communities and consequently crop yield. This study was aimed to evaluate the impact of technical-grade fungicide tebuconazole on plant growth promoting potentials of tebuconazole-tolerant Rhizobium isolate MRP1. The performance of the isolate MRP1-inoculated pea plants grown in tebuconazole treated soils was also assessed. Generally, the three concentrations [100 (recommended dose), 200 and 300 μg kg⁻¹ soil] of tebuconazole when used alone, adversely affected the growth, symbiosis, grain yield and nutrient uptake by pea plants. Concentration dependent phytotoxicity of tebuconazole was observed for all the measured parameters. On the contrary, fungicide tolerant Rhizobium sp. MRP1 in the presence of fungicide increased the measured parameters at all tested concentrations. As an example, when inoculant MRP1 was also used with 300 μg tebuconazole kg⁻¹ soil, it substantially increased the root nitrogen, shoot nitrogen, root P, shoot P, seed yield and grain protein by 20, 19, 50, 31, 15 and 7%, respectively, when compared with uninoculated plants grown in fungicide-treated soils. The study suggests that the plant growth promoting Rhizobium sp. MRP1 can be used as bacterial inoculant to increase the production of pea in soils polluted with fungicides.     

Abundance–occupancy and abundance–body mass relationships of small mammals in a mountainous landscape

Context

Mountainous landscapes are characterized by strong spatial heterogeneity, leading to increasing geographical isolation and decreasing area with elevation. Consequently, the colonization rate decreases from the low to high elevation zone, while the extinction rate shows the opposite.

Objectives

Due to such changes, we test whether (1) species occur at a declining number of sites (mountains) and have a less positive abundance–occupancy relationship (AOR) in a higher elevation zone; (2) a lower proportion of rare large-bodied species (less resistant to extinction) and a more positive abundance–body mass relationship (ABR) emerge in a higher elevation zone.

Methods

Using the data of small mammals from 20 elevational gradients in the Mountainous Region of Southwest China, we compared the AORs and ABRs among the low, middle and high elevation zones. The AOR and ABR were fitted with linear and polynomial regression models. We compared endemic ratios among the different elevation zones.

Results

The AOR was best characterized by a linear model and positive in all elevation zones. Its slope decreased from the low to high elevation zone. The quadratic model performed the best in fitting the ABR in each zone. When fitted with linear models, both the R² and slope of the ABR increased towards the high elevation zone. The endemic ratios were significantly higher in the middle and high elevation zones.

Conclusions

Both the AOR and ABR in mountainous landscapes are strongly elevation-dependent. The increasing geographical isolation and decreasing area with elevation can have a high impact on macroecological patterns and processes.

     

The use of a GIS in assessing the impacts of sea level rise on nature conservation along the Dutch coast: 1990–2090

The Dutch coastline is expected to change considerably during the next 100 years. Erosion will prevail, although accretion will occur locally.To establish a new policy for coastal defence management an integrated policy analysis study was performed. Major dune functions (nature conservation, recreation, public drinking water supply, housing and industry) have been inventarized by using a Geographic Information System.This study reports on the part of the analysis which takes nature conservation interests into account. Evaluation of nature interests has been based on the criteria: rarity and diversity of plant species c.q. vegetation types, succession stage, and completeness. This results in a classification of nature into five classes, based on abiotic as well as biotic characteristics of the landscape. Losses of nature interests were studied due to: loss of land because of shoreline retreat, to remodelling of the foredune ridge, and to changes of the dune groundwater level.An evaluation is given of the methods used to assess ecological impacts.Ideas are presented for further research on the prediction of ecological impacts and on coastline management which combines traditional coastal defence with nature conservation.     

Effects of the application of ‘cash’ on the growth,fruit yield,and nutrient status of tomato (Solanum lycopersicum L.) grown in reclaimed saline soil

Summary

The effects of ‘cash’ [a novel 2:5:1 (w/w/w) mixture of calcium sulphate, ground sunflower heads, and humic acid] as a soil amendment on the growth, fruit yield, and leaf nutrient status of tomato (Solanum lycopersicum L.) grown on reclaimed saline soil (EC = 9.4 dS m^–1) were investigated. A glasshouse experiment was performed in a completely randomised design with six treatments (0, 10, 20, 30, 40, or 50 g ‘cash’ kg^–1 soil) each with four replicates. The results indicated that ‘cash’ increased both the shoot dry weight (DW) plant^–1 and the root DW plant^–1, the free proline contents of leaves, and leaf chlorophyll contents. The use of ‘cash’ also increased the number of fruit plant^–1, fruit yield pot^–1, and fruit vitamin C contents, although total soluble solids (TSS) contents were not affected. The contents of nitrogen (N), potassium (K), and calcium (Ca), and the Ca:Na ratios of leaf tissues increased with all application rates of ‘cash’. Leaf phosphorus (P) contents showed no response to any applied level of ‘cash’. Leaf sodium (Na) contents declined gradually with an increase in the level of ‘cash’ applied to the soil. We concluded that ‘cash’ has a pronounced positive effect on the growth and fruit yield of tomato plants grown in reclaimed saline soil. ‘Cash’ therefore has the potential to be used as a soil amendment for vegetable crops such as tomato to overcome the adverse effects of salinity in newly-reclaimed soils.     

Thresholds of landscape change: a new tool to manage green infrastructure and social–economic development

An understanding of how and where a landscape can be improved with green infrastructure is important for the development of land-use policies. However, it is still a big challenge to manage landscapes due to a lack of condition-based diagnosis and consistent conflicts with social–economic interests. The purpose of this study is to identify the thresholds of landscape change and to explore new reference lines of policy decision-making for balancing social–economic development and green infrastructure management. Five different landscape types and their thresholds of landscape change were identified using parametric and piecewise linear models in the district of Haidian, Beijing, PR China, and their changes and social–economic drivers were analyzed with principal component analysis and a stepwise linear regression model for the time period 1991–2010. It is shown that different thresholds of different landscapes do exist and can be identified by the area of their key elements of green infrastructure. Integrating these thresholds into a social–economic context, it is shown where and how social–economic variables can be manipulated quantitatively to achieve development targets with respect to green infrastructure in individual towns and the entire district. Green infrastructure can be managed by changing just a small proportion of social–economic investment. This paper provides a useful tool to achieve a sustainable development by balancing green infrastructure with social–economic interests.     

Effects of farmyard manure on soil organic carbon stock,the pattern of fertility build-up,and plant growth in ‘Mallika’ mango (Mangifera indica L.)

Summary

A field experiment was conducted on mango (Mangifera indica L.) to study the effects of farmyard manure (FYM) on soil organic carbon (SOC) content and on the build-up of fertility over three growing seasons (2006–2007, 2007–2008, and 2008–2009) at Bhubaneswar, Orissa, India. Five treatments, including the application of FYM at 5.0 kg plant^–1, 7.5 kg plant^–1, or 10.0 kg plant^–1, the recommended dose of a chemical fertiliser (RDF; N:P:K=75:20:60), and a non-fertilised control, were evaluated in a randomised block design. The results indicated that SOC contents increased significantly in the FYM-treated plots. The highest SOC density (554 g m^–2) and stock (5.55 Mg ha^–1) were recorded in the treatment with FYM at 10 kg plant^–1. The rate of increase in SOC density was highest (193 g m^–2 year^–1) with FYM at 10 kg plant^–1, and lowest in the untreated control (15 g m^–2 year^–1). Regular addition of FYM had a positive effect on the build-up of soil fertility. However, the greatest increases in soil N, P, K contents were in the 7.5 kg plant^–1 FYM treatment (means = 319.9, 19.1, and 270.8 kg ha^–1 for N, P, K, respectively). Farmyard manure significantly influenced the growth parameters of mango trees over the three seasons. There was a positive and linear relationship between increasing rates of application of FYM and trunk cross-sectional area.     

Acclimatization of Codiaeum variegatum L. ‘Aucubaefolia’ to changes in photon flux density. II. Net photosynthesis,dark respiration and chlorophyll content

Summary

Net photosynthesis, dark respiration and chlorophyll content were studied in plants of Codiaeum when transferred” from a glasshouse to photon flux densities 6, 18, or 66 μE m^?2s^?1 in growth rooms. The diurnal patterns of daily net photosynthesis and dark respiration were recorded during the first 12 days and 3, 7, 11 and 19 weeks after the transfer. Only plants transferred to 66 μE m^?2s^?1 reached a positive total net photosynthesis per day within the first 12 days. This was achieved only between the 12th day and 3 weeks when transferred to 6 μE m^?2s^?1. The rate of net photosynthesis increased until the 11th week and then declined, while dark respiration decreased until the 11th week and then increased. The diurnal patterns of net photosynthesis and dark respiration changed significantly with time of acclimatization. The rate of change in daily net photosynthesis and dark respiration was highest in the plants transferred to the lowest photon flux density. Changes in chlorophyll content of the leaves were similar to those in net photosynthesis.     

Critical upscaling. How citizens’ initiatives can contribute to a transition in governance and quality of urban greenspace

Studies show that municipalities often develop a type of urban greenspace that is rather uniform in its shape and use. Citizens’ initiatives develop different types of greenspace. This article uses concepts from transition studies and identifies what happens during a citizens’ initiative in urban greenspace in the Netherlands in terms of transition of municipal management and development and how these initiatives can lead to a change of practices of the municipality. A single, qualitative study of Diepenheim Inside-Out-Forest in the Netherlands as a critical case is presented, based on 8 semi-open interviews. The study gives insight in how the municipality has changed in relation to this one case, and how such transition of ‘regime’ takes place. The study explains how the different benefits that arise in a greenspace development and management initiative relate to the ‘critical knowledge’ and ‘situated knowledge’ of the actors involved and that the quality of urban greenspace is very much the result of that knowledge. ‘Fit and conform’ and ‘stretch and transform’ are usable strategies for the empowerment of such initiatives through sharing of resources, and policy advocacy by ‘critical niche’ innovators.‘Regime’ is not uniquely a feature of the local state but also of market parties and citizens themselves with their own values and routines. A change among all parties seems needed if greenspace is to be developed, managed and used differently.     

Induction of resistance to Penicillium digitatum and chilling injury in ‘Star Ruby’ grapefruit by a short hot-water rinse and brushing treatment

Summary

Postharvest heat treatments have been used for many years as alternatives to chemical control of fungal diseases and insect infestation of fruits and vegetables. In this study, the effects of a new hot-water brushing (HWB) treatment on the resistance of red grapefruit (C. paradisi cv. Star Ruby) to green mould decay caused by Penicillium digitatum (Pers.: Fr.) Sacc. and on the development of chilling injury (CI) symptoms during cold storage were examined. The HWB treatment comprises rinsing hot water on the fruits as they move along a belt of brush rollers. A twenty second HWB treatment at 59 or 62° reduced decay, after arti®cial inoculation of wounded fruit, by 52 or 70%, respectively, compared with control unwashed fruit, whereas rinsing and brushing the fruit with tap water (~20°) or with hot water at 53 or 56°, were ineffective. HWB treatments applied 1–3 d prior to inoculation were most effective in enhancing the disease resistance of fruit, but were much less effective when the fruit were inoculated on the same day or 7.d later. HWB treatments at 59 or 62° for 20.s also significantly reduced the CI index and the percentage of fruit displaying CI symptoms by 42 and 58%, respectively, after six weeks’ storage at 2° and an additional week at 20°. Furthermore, HWB treatments cleaned the fruit and improved its general appearance without causing any surface damage, and did not influence fruit weight loss, percentage of total soluble solids (TSS) in the juice, juice acidity or fruit colour.     

Using a landscape ecological perspective to analyze regime shifts in social–ecological systems: a case study on grassland degradation of the Tibetan Plateau

Landscape Ecology - Landscape ecology thinking and social–ecological system (SES) thinking investigate human–environment relationships from the perspective of ‘space’ and...     

Influence of soil hydrothermal environment,irrigation regime,and different mulches on the growth and fruit quality of strawberry (Fragaria × ananassa L.) plants in a sub-temperate climate

Summary

The present study was undertaken over two consecutive years under sub-temperate climatic conditions in the mid-hill region of Himachal Pradesh (30°52'N, 77°11'E; 1,175 m asl) on loamy sand Inceptisols. The aim was to investigate the effects of irrigation and mulch material on the growth, flowering, fruiting behaviour, relative leaf water content (RLWC), yield, and quality of strawberry (Fragaria × ananassa L. ‘Chandler’). The drip irrigation treatments included irrigation with 100, 80, or 60% (coded 1.0, 0.8, and 0.6 V) of the total water requirement. Both mulches increased the minimum soil temperature to a depth of 5 cm. The drip and surface irrigation treatments raised the minimum soil temperature by 3.0º – 5.4ºC, and lowered the maximum temperature by 2.2º – 5.8ºC compared to the rainfed control. Hay mulch was more effective in raising the minimum temperature and lowering the maximum soil temperature than black polyethylene mulch. Moisture conservation increased by 2.8 – 12.8% under the black polyethylene mulch compared to the no-mulch treatment. Drip and surface irrigation methods, as well as mulching, were found to be effective for enhancing the growth, irrigation water use efficiency (IWUE), fruit yield, and quality of strawberry plants. However, the number of crowns per plant, the percentage berry set, the RLWC, root length density (RLD), and fruit yield were highest under treatment M3I3 [i.e., black polyethylene mulch plus drip irrigation (1.0 V)] by 565.5%, 94.5%, 32.8%, 394.5%, and 549.6%,respectively, compared to the no mulch plus rainfed control. The maximum IWUE values for plant biomass [1.39 metric tonnes (MT) ha^–1 m^–1] and fruit yield (2.79 MT ha^–1 m^–1) were recorded under treatment M3I5 [i.e., black polyethylene mulch plus drip irrigation (0.6 V)]; whereas, the lowest values (0.39 and 0.68 MT ha^–1 m^–1, respectively) were observed using treatment M1I2 (i.e., without mulch, plus surface irrigation). Fruit size, weight, sugar content, and anthocyanin content increased significantly under treatment M2I3 [i.e., hay mulch, plus drip irrigation (1.0 V)] compared with all other treatments. Total soluble solids (TSS) contents and total acidity (TA) were highest under treatment M1I1 (i.e., no mulch, plus rainfed). A linear regression model could describe the variations in quality parameters of strawberry plants grown under sub-temperate climatic conditions. Root density was found to be the best indicator with which estimate fruit quality.     

Using an individual-based model to examine the roles of habitat fragmentation and behavior on predator–prey relationships in seagrass landscapes

Seagrasses, which form critical subtidal habitats for marine organisms worldwide, are fragmented via natural processes but are increasingly being fragmented and degraded by boating, fishing, and coastal development. We constructed an individual-based model to test how habitat fragmentation and loss influenced predator–prey interactions and cohort size for a group of settling juvenile blue crabs (Callinectes sapidus Rathbun) in seagrass landscapes. Using results from field studies suggesting that strong top-down processes influence the relationship between cannibalistic blue crab populations and seagrass landscape structure, we constructed a model in which prey (juvenile blue crabs) are eaten by mesopredators (larger blue crabs) which in turn are eaten by top-level predators (e.g., large fishes). In our model, we varied the following parameters within four increasingly fragmented seagrass landscapes to test for their relative effects on cohort size: juvenile blue crab (prey) predator avoidance response, hunting ability of mesopredators and predators, the presence of a top-level predator, and prey settlement routines. Generally, prey cohort size was maximized in the presence of top-level predators and when mesopredators and predators exhibited random searching behavior vs. directed hunting. Cohort size for stationary (tethered) prey was maximized in fragmented landscapes, which corresponds to results from field experiments, whereas mobile prey able to detect and avoid predators had higher survival in continuous landscapes. Prey settlement patterns had relatively small influences on cohort size. We conclude that the effects of seagrass fragmentation and loss on organisms such as blue crabs will depend heavily on behaviors of prey and predatory organisms and how these behaviors change with landscape structure.