Ecoinformatics Reveals Effects of Crop Rotational Histories on Cotton Yield

Crop rotation has been practiced for centuries in an effort to improve agricultural yield. However, the directions, magnitudes, and mechanisms of the yield effects of various crop rotations remain poorly understood in many systems. In order to better understand how crop rotation influences cotton yield, we used hierarchical Bayesian models to analyze a large ecoinformatics database consisting of records of commercial cotton crops grown in California''s San Joaquin Valley. We identified several crops that, when grown in a field the year before a cotton crop, were associated with increased or decreased cotton yield. Furthermore, there was a negative association between the effect of the prior year''s crop on June densities of the pest Lygus hesperus and the effect of the prior year''s crop on cotton yield. This suggested that some crops may enhance L. hesperus densities in the surrounding agricultural landscape, because residual L. hesperus populations from the previous year cannot continuously inhabit a focal field and attack a subsequent cotton crop. In addition, we found that cotton yield declined approximately 2.4% for each additional year in which cotton was grown consecutively in a field prior to the focal cotton crop. Because L. hesperus is quite mobile, the effects of crop rotation on L. hesperus would likely not be revealed by small plot experimentation. These results provide an example of how ecoinformatics datasets, which capture the true spatial scale of commercial agriculture, can be used to enhance agricultural productivity.     

Influence of the surrounding landscape on crop colonization by a polyphagous insect pest

Landscape composition plays an important, but poorly understood, role in the population dynamics of agricultural pest species with broad host ranges including both crops and weeds. One such pest, the generalist plant bug Lygus hesperus Knight (Hemiptera: Miridae), is a key cotton pest that feeds on various hosts differing in quality in California's San Joaquin Valley (USA). We investigated the effects of 15 common crops and uncultivated agricultural land on L. hesperus populations, by correlating the densities of L. hesperus in focal cotton fields with the areas of the 16 crops in surrounding rings. Insect counts were provided by private pest‐control advisors, and spatial data were obtained from Kern County records. We first calculated Spearman's partial correlation coefficients on an annual basis for each crop separately, and then performed a meta‐analysis of these correlations across years to describe the overall effect of a particular crop on L. hesperus after the effects of the 15 other crops are removed. Consistent with studies conducted in other areas, L. hesperus density was positively correlated with safflower, and negatively with cotton. Lygus hesperus density was also correlated with several other crops that are often not considered in pest management, including grape, oat, and onion (positive correlations), and almond, pistachio, and potato (negative correlations). Lygus hesperus density was also found to be negatively correlated with alfalfa and positively correlated with uncultivated habitats, a relationship that receives mixed support in the literature. Several other crops tested were not significantly correlated with L. hesperus densities in focal cotton fields, suggesting a neutral role for them in L. hesperus dynamics. The improved understanding of the effects of a greater variety of crops on L. hesperus population dynamics will be useful in the design of agricultural landscapes for enhanced management of this important polyphagous pest.     

Stage-specific predation on Lygus hesperus affects its population stage structure

Evidence suggests that prey can vary in their susceptibility to predation depending on their developmental stage. Stage‐dependent predation is of particular importance to integrated pest management, because it is often a particular developmental stage of a pest that causes the majority of the damage to the crop. An understanding of stage‐dependent biocontrol is therefore important for predicting the ultimate impact of herbivore populations. In this article, we addressed this issue by focusing on the stage structure of Lygus hesperus Knight (Heteroptera: Miridae) populations in cotton as related to the density of a specific generalist predator. We first demonstrated in a field experiment that Geocoris spp. adults suppressed L. hesperus eggs and/or early instars (first through third), but did not suppress fourth through fifth instars or adult L. hesperus. We then demonstrated that this stage‐specific predation translated into season‐long shifts in the stage structure of L. hesperus populations in cotton fields. Using weekly sweep counts across 21 separate cotton fields, we found a negative correlation between season‐long Geocoris spp. densities and season‐long densities of L. hesperus nymphs. In contrast, there was no such correlation between Geocoris spp. and L. hesperus adults. Taken together, these results suggest that Geocoris spp. predators influence the stage structure of L. hesperus populations in cotton, which in turn has the potential to affect patterns of cotton square damage and sampling bias when monitoring L. hesperus populations.     

A GIS-based approach for areawide pest management: the scales of Lygus hesperus movements to cotton from alfalfa, weeds, and cotton

Understanding the effect of cropping patterns on population dynamics, dispersal, and habitat selection of insect pests has been an unresolved challenge. Here, we studied the western tarnished plant bug, Lygus hesperus (Knight) (Heteroptera: Miridae), in cotton during early summer in central Arizona. We used a general approach based on global positioning system (GPS) and geographic information system (GIS) technologies combined with spatial statistics to assess the maximum distance at which forage and seed alfalfa, fallow fields with weeds, and cotton affect L. hesperus population density. Using a set of 50 cotton fields as focal fields, we found that forage and seed alfalfa as well as weeds acted as L. hesperus sources for these cotton fields. The source effect did not extend beyond 375, 500, and 1500 m for forage alfalfa, weeds, and seed alfalfa, respectively. Conversely, cotton fields acted as L. hesperus sinks, but this effect did not extend further than 750 m from the focal cotton fields. These findings suggest that specific spatial arrangements of these field types could reduce L. hesperus damage to cotton. The spatially explicit approach used here provides a direct evaluation of the effects of agroecosystem heterogeneity on pest population dynamics, dispersal, and habitat selection, which is a significant asset for the development and improvement of areawide pest management.     

Ecoinformatics for integrated pest management: expanding the applied insect ecologist's tool-kit

Experimentation has been the cornerstone of much of integrated pest management (IPM) research. Here, we aim to open a discussion on the possible merits of expanding the use of observational studies, and in particular the use of data from farmers or private pest management consultants in "ecoinformatics" studies, as tools that might complement traditional, experimental research. The manifold advantages of experimentation are widely appreciated: experiments provide definitive inferences regarding causal relationships between key variables, can produce uniform and high-quality data sets, and are highly flexible in the treatments that can be evaluated. Perhaps less widely considered, however, are the possible disadvantages of experimental research. Using the yield-impact study to focus the discussion, we address some reasons why observational or ecoinformatics approaches might be attractive as complements to experimentation. A survey of the literature suggests that many contemporary yield-impact studies lack sufficient statistical power to resolve the small, but economically important, effects on crop yield that shape pest management decision-making by farmers. Ecoinformatics-based data sets can be substantially larger than experimental data sets and therefore hold out the promise of enhanced power. Ecoinformatics approaches also address problems at the spatial and temporal scales at which farming is conducted, can achieve higher levels of "external validity," and can allow researchers to efficiently screen many variables during the initial, exploratory phases of research projects. Experimental, observational, and ecoinformatics-based approaches may, if used together, provide more efficient solutions to problems in pest management than can any single approach, used in isolation.     

Stage-dependent feeding behavior by western tarnished plant bugs influences flower bud abscission in cotton

Unexplained variability in the relationship between the number of herbivores in a field and the amount of crop damage can arise if there is a large amount of variation among herbivore individuals in the amount of feeding damage each generates. In California, populations of the western tarnished plant bug, Lygus hesperus Knight (Heteroptera: Miridae), produce highly variable levels of damage to cotton plants (Gossypium hirsutum L.) (Malvaceae), even when found at low densities. Because L. hesperus populations are also highly variable in their overall stage structure, we hypothesize that differences in crop damage might result from varying impact by each L. hesperus stage on cotton flower buds (termed squares). Laboratory measurements of L. hesperus mouth‐parts and distance to anther sacs, a preferred feeding site, revealed that 1st?3rd instar L. hesperus nymphs will not be able to feed on anther sacs of larger squares (over 8 mm in length) but will be able to feed on squares that are most sensitive to L. hesperus damage (<7 mm). Because even the 1st instars can feed on the most sensitive ‘pinhead’ squares, size constraints do not rule out damaging effects from the youngest L. hesperus. Laboratory observations revealed that later developmental stages, and adults, spend more time feeding on cotton squares relative to 2nd and 3rd instars. In addition, a field experiment revealed no effect of 2nd instars on square retention (relative to control cages) but did reveal a significant decrease in square retention generated by adult L. hesperus (4th instar L. hesperus resulted in an intermediate level of square retention). In a final study we sampled L. hesperus stage structure and density across 38 cotton fields. Multiple regression revealed that the densities of 1st?3rd instars of L. hesperus are not correlated with anther sac damage or square retention. However, in 2 years 4th and 5th instars were positively correlated with anther sac damage and negatively correlated with square retention. In the a third year, adult L. hesperus showed correlations in the same direction, across fields and across sites within fields. Overall, these results suggest that the adults and the largest nymphs of L. hesperus (4th and 5th instars) are particularly damaging to cotton squares, with the 1st?3rd instars of L. hesperus causing little damage to plants.     

State-dependent sampling bias in insects: implications for monitoring western tarnished plant bugs

Insect populations vary in the proportion of individuals exhibiting a particular ‘state’ (e.g., developmental stage, sex, egg load, or nutritional status). Because an insect's developmental state often determines both its size and its behavior, it is likely that this will also affect the probability of being sampled. We propose that a comprehensive approach to pest management must consider the interaction between the structure of a pest population (i.e., the relative number of individuals in each state) and any state‐dependent sampling bias. To illustrate the usefulness of this method we sampled populations of the western tarnished pest bug, Lygus hesperus Knight (Heteroptera: Miridae), in cotton fields. Our sampling technique utilized large cages to measure the absolute densities of each L. hesperus stage and adult sex within a population. This technique allowed us to document a wide range of absolute stage structures and sex ratios across 10 L. hesperus populations in California. Using a combination of cage samples and sweep net samples, we quantified the state‐dependent sampling bias by calculating the efficiency of sweep sampling as a function of L. hesperus developmental stage and sex. We found that the efficiency of sweep nets increased steadily with each successive developmental stage (i.e., nymphal instar) of L. hesperus. We also found that sweep nets are slightly more efficient in capturing male vs. female L. hesperus adults. Since other studies have documented that the stage and sex of L. hesperus can affect feeding impact on cotton flower buds, our results suggest that accurate predictions of Lygus damage will need to incorporate stage and sex‐dependent sampling biases.     

Identifying existing management practices in the control of Striga asiatica within rice–maize systems in mid‐west Madagascar

Infestations by the parasitic weed genus Striga result in significant losses to cereal crop yields across sub‐Saharan Africa. The problem disproportionately affects subsistence farmers who frequently lack access to novel technologies. Effective Striga management therefore requires the development of strategies utilizing existing cultural management practices. We report a multiyear, landscape‐scale monitoring project for Striga asiatica in the mid‐west of Madagascar, undertaken over 2019–2020 with the aims of examining cultural, climatic, and edaphic factors currently driving abundance and distribution. Long‐distance transects were established across the middle‐west region of Madagascar, over which S. asiatica abundance in fields was estimated. Analysis of the data highlights the importance of crop variety and legumes in driving Striga density. Moreover, the dataset revealed significant effect of precipitation seasonality, mean temperature, and altitude in determining abundance. A composite management index indicated the effect of a range of cultural practices on changes in Striga abundance. The findings support the assertion that single measures are not sufficient for the effective, long‐term management of Striga. Furthermore, the composite score has potential as a significant guide of integrated Striga management beyond the geographic range of this study.     

Population analysis of white grubs (Coleoptera: Melolonthidae) throughout the Brazilian Pampa biome

The replacement of natural grassland by cultivated areas might favor the increase in abundance of some root-feeding species such as the white grubs, which may become a constraint for field crop production. This research aimed to assay the population density and geographical distribution of white grubs pest and other species in natural grassland and cultivated areas throughout the Brazilian Pampa biome. White grubs were sampled in 18 locations in both landscape use types and identified. Population density (number of larvae m^?2) was calculated for each recorded species and sorted within two groups (pest species and other species), compared between natural grasslands and cultivated areas, as well as among locations. A dendrogram to evaluate species similarity among locations was built based on combined data obtained from both landscape use types throughout the region. In total, 31 species were found in the Brazilian Pampa, and four of them are considered as crop pests: Diloboderus abderus (Sturm, 1826), Euetheola humilis (Burmeister, 1847), Lyogenys fusca (Blanchard, 1830), and Phyllophaga triticophaga Morón & Salvadori, 1998. The average population density of pest species in cultivated areas was less than five larvae m^?2, at most of locations. Some species had a wide geographical distribution (e.g. D. abderus and Cyclocephala modesta Burmeister), while other melolontids occurred at only one location. The knowledge of which white grub species are present in a field and its population densities assist farmers to take proper management decisions.     

Yield Reduction and Root Damage to Cotton Induced by Belonolaimus longicaudatus

Sting nematode (Belonolaimus longicaudatus) is recognized as a pathogen of cotton (Gossypium hirsutum), but the expected damage from a given population density of this nematode has not been determined. The objective of this study was to quantify the effects of increasing initial population densities (Pi) of B. longicaudatus on cotton yield and root mass. In a field plot study, nematicide application and cropping history were used to obtain a wide range of Pi values. Cotton yields were regressed on Pi density of B. longicaudatus to quantify yield losses in the field. In controlled environmental chambers, cotton was grown in soil infested with increasing Pi''s of B. longicaudatus. After 40 days, root systems were collected, scanned on a desktop scanner, and root lengths were measured. Root lengths were regressed on inoculation density of B. longicaudatus to quantify reductions in the root systems. In the field, high Pi''s (>100 nematodes/130 cm³ of soil) reduced yields to near zero. In controlled environmental chamber studies, as few as 10 B. longicaudatus/130 cm³ of soil caused a 39% reduction in fine cotton roots, and 60 B. longicaudatus/130 cm³ of soil caused a 70% reduction. These results suggest that B. longicaudatus can cause significant damage to cotton at low population densities, whereas at higher densities crop failure can result.     

Species composition and population dynamics of some arthropod pests in cotton fields of irrigated and semi-arid regions of Punjab,Pakistan

The present study aimed to record seasonal dynamics and diversity of different insect pests in cotton agroecosystems. Two well-known cotton growing districts i.e. district Layyah and Vehari were selected for the study from the cotton belt of Punjab, Pakistan. Sampling was done bi-weekly for two consecutive years from July to October during 2018 and 2019. Sweep netting, visual counting, hand picking, wet finger method, beat sheets, aspirator and pitfall trapping methods were used for collection. Shannon-Wiener and Simpson indices were used to compute diversity while Menhinick and Margalef indices were used for the estimation of species richness. A total of 94,343 individuals representing 43 species, 40 genera, 28 families and 6 orders were recorded. Family Aleyrodidae dominated over other pest families. Bemisia tabaci (Gennadius, 1889) of family Aleyrodidae was the dominant species with 39.68% share among all pest species. Estimated species richness of all arthropod pest species belonging to both districts were about 94%. The densities of pests fluctuated with time. The peaks of sucking pest densities were observed in July and August while densities of chewing pests peaked in late September or early October each year. Population densities of jassids Amrasca biguttula (Ishida), armyworm Spodoptera litura (Fabricius), and pink bollworm Pectinophora gossypiella (Saunders), showed strong negative correlation with temperature, humidity and rainfall while thrips population density showed positive correlation with these parameters. Minor loss due to pests are acceptable everywhere, but it is only possible when their populations remain below their economic threshold levels. Present study will aid to design pest management strategies in cotton growing areas round the globe.     

Adjacent Habitat Influence on Stink Bug (Hemiptera: Pentatomidae) Densities and the Associated Damage at Field Corn and Soybean Edges

The local dispersal of polyphagous, mobile insects within agricultural systems impacts pest management. In the mid-Atlantic region of the United States, stink bugs, especially the invasive Halyomorpha halys (Stål 1855), contribute to economic losses across a range of cropping systems. Here, we characterized the density of stink bugs along the field edges of field corn and soybean at different study sites. Specifically, we examined the influence of adjacent managed and natural habitats on the density of stink bugs in corn and soybean fields at different distances along transects from the field edge. We also quantified damage to corn grain, and to soybean pods and seeds, and measured yield in relation to the observed stink bug densities at different distances from field edge. Highest density of stink bugs was limited to the edge of both corn and soybean fields. Fields adjacent to wooded, crop and building habitats harbored higher densities of stink bugs than those adjacent to open habitats. Damage to corn kernels and to soybean pods and seeds increased with stink bug density in plots and was highest at the field edges. Stink bug density was also negatively associated with yield per plant in soybean. The spatial pattern of stink bugs in both corn and soybeans, with significant edge effects, suggests the use of pest management strategies for crop placement in the landscape, as well as spatially targeted pest suppression within fields.     

Combining intercropping with semiochemical releases: optimization of alternative control of Sitobion avenae in wheat crops in China

Overreliance on pesticides has large environmental and human health costs that compel researchers and farmers to seek alternative management tactics for crop pests. For insect pests, increasing crop species diversity via intercropping and using semiochemicals to alter local arthropod populations have separately proven effective at reducing pest densities. Here, we combine these two tactics in an effort to gain better control of Sitobion avenae (Fabricius) (Hemiptera: Aphididae), the English grain aphid, a major pest of cereal production worldwide. We conducted field experiments over 2 years testing the effectiveness of combining intercropping of wheat and oilseed rape with release of methyl salicylate (MeSA). We found that maximum and mean aphid densities were highest in wheat monocultures, significantly lower in intercropped plots and MeSA plots, and lowest when intercropping and MeSA release were combined by obtaining highest densities of predatory lady beetles and parasitoids rates. Importantly, grain yield and quality showed a similar pattern: they were highest for combined intercropped/MeSA plots, intermediate in plots with intercropping or MeSA alone, and lowest in control monoculture plots. Our results suggest that combining these two tactics holds significant promise for improved management of aphid populations and emphasize the need to integrate alternative pest control approaches to optimize sustainable insect pest management.     

Maximizing the Potential of Cropping Systems for Nematode Management

Quantitative techniques were used to analyze and determine optimal potential profitability of 3-year rotations of cotton, Gossypium hirsutum cv. Coker 315, and soybean, Glycine max cv. Centennial, with increasing population densities of Hoplolaimus columbus. Data collected from naturally infested on-farm research plots were combined with economic information to construct a microcomputer spreadsheet analysis of the cropping system. Nonlinear mathematical functions were fitted to field data to represent damage functions and population dynamic curves. Maximum yield losses due to H. columbus were estimated to be 20% on cotton and 42% on soybean. Maximum at-harvest population densities were calculated to be 182/100 cm³ soil for cotton and 149/100 cm³ soil for soybean. Projected net incomes ranged from a $17.74/ha net loss for the soybean-cotton-soybean sequence to a net profit of $46.80/ha for the cotton-soybean-cotton sequence. The relative profitability of various rotations changed as nematode densities increased, indicating economic thresholds for recommending alternative crop sequences. The utility and power of quantitative optimization was demonstrated for comparisons of rotations under different economic assumptions and with other management alternatives.     

Impact of Cotton Production Systems on Management of Hoplolaimus columbus

The effectiveness of selected cultural practices in managing the Columbia lance nematode, Hoplolaimus columbus, on cotton was evaluated in experiments in growers'' infested fields. The effects of planting date, cotton cultivar, treatment with the growth regulator mepiquat chloride, and destruction of cotton-root systems after harvest on cotton-lint yield and population densities of H. columbus were studied. The yield of cotton cultivar Deltapine 50 was negatively related (P = 0.054) to initial population density of H. columbus whereas the yield of Deltapine 90 was not affected by preplant density of this nematode, indicating tolerance in Deltapine 90. Reproduction of this nematode did not differ on the two cultivars. Planting date and treatment with the growth regulator mepiquat chloride did not influence cotton yield in a consistent manner. Application of mepiquat chloride suppressed (P ≤ 0.05) numbers of Columbia lance nematode, although there was an interaction (P ≤ 0.05) with cultivar and year. Early vs. late destruction of cotton-root systems did not impact population densities of this nematode either year, and had no impact on the subsequent cotton crop. The nematicide fenamiphos increased (P ≤ 0.03) cotton yield when H. columbus numbers exceeded the damage threshold.     

Artificial miRNA-mediated silencing of ecdysone receptor (EcR) affects larval development and oogenesis in Helicoverpa armigera

The insect pests are real threat to farmers as they affect the crop yield to a great extent. The use of chemical pesticides for insect pest control has always been a matter of concern as they pollute the environment and are also harmful for human health. Bt (Bacillus thuringensis) technology helped the farmers to get rid of the insect pests, but experienced a major drawback due to the evolution of insects gaining resistance towards these toxins. Hence, alternative strategies are high on demand to control insect pests. RNA-based gene silencing is emerging as a potential tool to tackle with this problem. In this study, we have shown the use of artificial microRNA (amiRNA) to specifically target the ecdysone receptor (EcR) gene of Helicoverpa armigera (cotton bollworm), which attacks several important crops like cotton, tomato chickpea, pigeon pea, etc and causes huge yield losses. Insect let-7a precursor miRNA (pre-miRNA) backbone was used to replace the native miRNA with that of amiRNA. The precursor backbone carrying the 21 nucleotide amiRNA sequence targeting HaEcR was cloned in bacterial L4440 vector for in vitro insect feeding experiments. Larvae fed with Escherichia coli expressing amiRNA-HaEcR showed a reduction in the expression of target gene as well as genes involved in the ecdysone signaling pathway downstream to EcR and exhibited mortality and developmental defects. Stem-loop RT-PCR revealed the presence of amiRNA in the insect larvae after feeding bacteria expressing amiRNA-HaEcR, which was otherwise absent in controls. We also found a significant drop in the reproduction potential (oogenesis) of moths which emerged from treated larvae as compared to control. These results demonstrate the successful use of an insect pre-miRNA backbone to express amiRNA for gene silencing studies in insects. The method is cost effective and can be exploited as an efficient and alternative tool for insect pest management.     

Crop Rotation and Races of Meloidogyne incognita in Cotton Root-knot Management

The influence o f various crop rotations and nematode inoculum levels on subsequent population densities of Meloidogyne incognita races 1 and 3 were studied in microplots. Ten different 3-year sequences o f cotton, corn, peanut, or soybean, all with cotton as the 3rd-year crop, were grown in microplots infested with each race. Cotton monoculture, two seasons o f corn, or cotton followed by corn resulted in high race 3 population densities and severe root galling on cotton the 3rd year. Peanut for 2 years preceding cotton most effectively decreased the race 3 population and root galls on cotton the 3rd year. Race 1 did not significantly influence cotton growth or yield at initial populations of up to 5,000 eggs/500 cm³ soil. At 5,000 eggs/500 cm³, cotton growth was suppressed by race 3 but yield was not affected.     

Effects of climate change on different geographical populations of the cotton bollworm Helicoverpa armigera (Lepidoptera,Noctuidae)

The effects of climate change on pest phenology and population size are highly variable. Understanding the impacts of localized climate change on pest distribution and phenology is helpful for improving integrated pest management strategies. Here, the population dynamics of cotton bollworms (Helicoverpa armigera) from Maigaiti County, south Xinjiang, and Shawan County, north Xinjiang, China, were analyzed using a 29‐year dataset at lower latitudes and a 23‐year dataset at higher latitudes to determine the effects of climate change on the population dynamics of H. armigera. The results showed that all generations of H. armigera at both sites showed increasing trends in population size with climate warming. Abrupt changes in phenology and population number occurred after abrupt temperature changes. Climate change had a greater effect on the phenology of H. armigera at higher latitudes than at lower latitudes and led to a greater increase in population size at lower latitudes than at higher latitudes; the temperature increase at higher latitudes will cause a greater increase in the adult moth population size in the future compared to that at lower latitudes; and abrupt changes in the phenology, temperature increase, and population size at lower latitudes occurred earlier than those at higher latitudes. Thus, it is necessary to develop sustainable management strategies for Helicoverpa armigera at an early stage.     

Supplemental releases of specialist parasitic wasps improve whitefly and psyllid control by <Emphasis Type="Italic">Dicyphus hesperus</Emphasis> in tomato

Dicyphus hesperus Knight (Heteroptera: Miridae) can contribute to the suppression of populations of Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae) and Bactericera cockerelli Sulcer (Hemiptera: Psyllidae) in tomato. Nevertheless, the remaining levels of these pests could still be too high for the crop to tolerate. We thus tested here whether the combination of D. hesperus with the specialist parasitoids Eretmocerus eremicus Rose & Zolnerowich (Hymenoptera: Aphelinidae) (whitefly) and Tamarixia triozae (psyllid) can result in better pest control compared with methods based exclusively on single-species releases in tomato. We conducted two simultaneous experiments in tomato (‘Whitefly’ and ‘Psyllid’ Experiment), where we compared the effectiveness against B. tabaci and B. cockerelli in cages receiving releases of the predator or the specialist parasitoid alone, or in combination. Although all natural enemies reduced pest levels when released separately, the combination of D. hesperus with E. eremicus and D. hesperus with T. triozae resulted in better whitefly and psyllid control, respectively, compared with the separate releases.