Impact of brown stink bug (Heteroptera: Pentatomidae) feeding on corn grain yield components and quality

Brown stink bug, Euschistus servus (Say) (Heteroptera: Pentatomidae), damage on developing corn, Zea mays L., ears was examined in 2005 and 2006 by using eight parameters related to its yield and kernel quality. Stink bug infestations were initiated when the corn plants were at tasseling (VT), mid-silking (R1), and blister (R2) stages by using zero, three, and six in 2005 or zero, one, two, and four bugs per ear in 2006, and maintained for 9 d. The percentage of discolored kernels was affected by stink bug number in both years, but not always affected by plant growth stage. The growth stage effect on the percentage of discolored kernels was significant in 2006, but not in 2005. The percentage of aborted kernels was affected by both stink bug number and plant growth stage in 2005 but not in 2006. Kernel weight was significantly reduced when three E. sercus adults were confined on a corn ear at stage VT or R1 for 9 d in 2005, whereas one or two adults per ear resulted in no kernel weight loss, but four E. servus adults did cause significant kernel weight loss at stage VT in 2006. Stink bug feeding injury at stage R2 did not affect kernel damage, ear weight or grain weight in either year. The infestation duration (9 or 18 d) was positively correlated to the percentage of discolored kernels but did not affect kernel or ear weight. Based on the regression equations between the kernel weight and stink bug number, the gain threshold or economic injury level should be 0.5 bugs per ear for 9 d at stage VT and less for stage R1. This information will be useful in developing management guidelines for stink bugs in field corn during ear formation and early grain filling stages.     

Toxicity of insecticides in a glass-vial bioassay to adult brown, green, and southern green stink bugs (Heteroptera: Pentatomidae)

Adult brown, Euschistus servus (Say); green, Acrosternum hilare (Say); and southern green, Nezara viridula (L.), stink bugs were collected from soybean, Glycine max (L.) Merr., in fall 2001 and 2002 near Stoneville, MS, and Eudora, AR. A glass-vial bioassay was used to determine LC50 values for the three species of stink bugs for the pyrethroids bifenthrin, cypermethrin, cyfluthrin, lambda-cyhalothrin, and permethrin, and the organophosphates acephate, dicrotophos, malathion, and methyl parathion. Results confirmed findings of other researchers that the brown stink bug was less susceptible to pyrethroid and organophosphate insecticides than were green and southern green stink bugs. The susceptibility of all three stink bug species to the insecticides tested was very similar at both test locations. The study established baseline insecticide mortality data from two locations in the mid-South for three stink bug species that are pests of soybean and cotton, Gossypium spp. Data from the tests are valuable for future use in studies on resistance and in resistance monitoring programs.     

Dynamics and seasonal abundance of stink bugs (Heteroptera: Pentatomidae) in a cotton-soybean ecosystem

Two soybean varieties (early-maturing group V and late-maturing group VII) and two cotton varieties (conventional and transgenic (Bt) were grown in adjacent replicated large field plots (approximately 0.1 ha each) at two locations for 3 yr. The dynamics and relative abundance of phytophagous stink bugs within these two crops were observed. The most abundant pentatomid species in both crops for all 3 yr were Nezara viridula (L.), Acrosternum hilare (Say), and Euschistus servus (Say). Several other species also were commonly collected. This is the first record of Mormidea lugens (F.) on soybean and E. quadrator Rolston, E. obscurus (Palisot), Holcostethus limbolarius (St?l), and Oebalus pugnax (F.) on cotton. Stink bugs began arriving in soybean when plant growth stages ranged from pod formation to full seed development. Peak numbers of these insects were found in soybean from the time of full-size seeds in the pods until early maturity. The bugs were first attracted to the earlier maturing cultivar (group V), where they remained until plants began to mature (R7). The pentatomids then moved to the later-maturing cultivar (group VII) as it reached full pod to full seed. Stink bugs began arriving in cotton from the time of the earliest flowers until after the first bolls formed. Peak numbers in cotton occurred during the time when all stages of developing bolls were present. Stink bug numbers were much greater in soybean than in cotton over all three seasons. This preference for soybean over cotton indicates the potential use of soybean as a trap crop for attracting stink bugs away from cotton. Additionally, the coordinated use of early- and late-maturing soybean cultivars as a trap crop could minimize the area requiring insecticides, as well as the number of insecticide applications to cotton.     

Injury to preflowering and flowering cotton by brown stink bug and southern green stink bug

The impact of brown stink bug, Euschistus servus (Say), and southern green stink bug, Nezara viridula (L.), injury was evaluated on preflowering and flowering cotton, Gossypium hirsutum L., plants in no-choice tests. Vegetative stage cotton seedlings and reproductive structures, including flower buds (square) and bolls, were infested with adults and/or nymphs of both species. There were no significant differences in height, height to node ratio, square retention, and flower initiation for cotton seedlings or plants with a match-head square between southern green stink bug adult- or brown stink bug adult-infested and noninfested treatments. Abscission for individual large squares (precandle) and multiple squares (medium and small square on the same sympodial branch) was not significantly different among infested and noninfested treatments for the following species and developmental stages: brown stink bug adults, southern green stink bug adults, and third and fourth to fifth instar southern green stink bug nymphs. In boll infestation studies, the relationship between boll maturity, expressed as heat units beyond anthesis, and boll growth, abscission, hard locked carpels, seedcotton yield, and seed germination was measured. Brown stink bug induced abscission in bolls that had accumulated > 0-350 heat units beyond anthesis. Boll growth and seedcotton yield was significantly lower for bolls infested with brown stink bug through 266.5 and 550 heat units beyond anthesis, respectively. The proportion of hard locked carpels per boll was significantly greater for the infested treatment in a cohort of bolls that accumulated from 51 to 400 heat units beyond anthesis. Seed germination in bolls infested with brown stink bug was significantly lower in bolls aged 101-600 heat units beyond anthesis.     

Biological control of soybean stink bugs by inoculative releases of Trissolcus basalis

Field studies of inoculative releases of Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) in early-maturing soybean, used as a trap crop, were conducted during four consecutive seasons to evaluate the parasitoid's potential as an IPM tactic for use against stink bug (Pentatomidae) species. Fifteen thousand adults of the parasitoid were released per hectare in the trap crop when the first stink bugs were detected in the experimental area. The stink bug population density was reduced by an average of 54% in the trap crop and by 58% in the main crop. The inoculative releases produced a reduction and delay in the stink bug population peak, mainly represented by Nezara viridula (L.), Piezodorus guildinii (Westwood) and Euschistus heros (Fabr.), which were held below economic threshold levels during the most critical stages of stink bug attack on soybean (pod and seed fill, R3–R6). As a result, seed quality was better in areas where T. basalis was released, demonstrating the efficacy of inoculative releases of this egg-parasitoid which could be an important component of the soybean IPM program in Brazil.     

Identification of Stink Bugs Using an Electronic Nose

Stink bugs are recognized as pests of several economically important crops, including cotton, soybean and a variety of tree fruits. The Cyranose 320 was used for the classified investigation of stink bug. Stink bugs including males and females of the southern green stink bugs, Nezara viridula, were collected from crop fields around College Station, TX. Results show that the released chemicals and chemical intensity are both critical factors, which determine the rate that the Cyranose 320 correctly identified the stink bugs. The Cyranose 320 shows significant potential in identifying stink bugs, and can classify stink bug samples by species and gender.     

Toxicity of Chromobacterium subtsugae to southern green stink bug (Heteroptera: Pentatomidae) and corn rootworm (Coleoptera: Chrysomelidae)

Diabrotica spp. (Coleoptera: Chrysomelidae) beetles and southern green stink bugs, Nezara viridula (L.) (Heteroptera: Pentatomidae), are pests on corn, Zea mays L., and soybean, Glycine max (L.) Merr., as well as on cucurbits. Control of these insects has depended on chemicals. An alternative to chemical control is the use of biologicals. Use of bacteria, fungi, viruses, pheromones, and metabolites to control these insects can potentially improve resistance management and reduce pesticide use. Other than Bacillus thuringiensis Berliner, few bacteria have been discovered that are lethal to either of these pests. Chromobacterium subtsugae Martin et al., a newly described bacterium that is known to be toxic to Colorado potato beetle, Leptinotarsa decemlineata (Say), larvae, was found to be toxic to both diabroticite adult beetles and southern green stink bug adults. In laboratory assays, toxins produced by these bacteria kill 80-100% of the adults of two species of diabroticite beetles, Diabrotica undecimpunctata howardi Barber and Diabrotica virgifera virgifera LeConte, and 100% of southern green stink bug adults within 6 d. For green stink bug, live bacteria were not needed for toxicity.     

Treatment thresholds for stink bugs (Hemiptera: Pentatomidae) in cotton

The green stink bug, Acrosternum hilare (Say), the southern green stink bug, Nezara viridula (L), and the brown stink bug, Euschistus servus (Say), were predominant phytophagous Pentatomidae detected during 1995-1997 in cotton in South Carolina. These species occurred in similar numbers in conventional and transgenic cotton 'NuCOTN33B', containing the gene for expression of CryIA(c) delta-endotoxin of Bacillus thuringiensis Berliner variety kurstaki. Adult stink bugs moved into cotton from wild and cultivated alternate hosts during July, and reproducing populations usually were detected in cotton from late July into September. Applications of either methyl parathion (0.56 kg [AI]/ha) directed for stink bugs or lambda-cyhalothrin (0.037 kg [AI]/ha) or cyfluthrin (0.056 kg [AI]/ha) for control of cotton bollworm, Helicoverpa zea (Boddie), provided effective control of pentatomids in NuCOTN33B or conventional 'DP5415' and increased yields compared with untreated plots. Fiber quality did not differ among treated or untreated plots of NuCOTN33B. The ground-cloth technique was used to estimate populations of stink bugs, and data indicated that treatment at one bug per 2 m of row adequately protected cotton from yield loss due to stink bug damage. Observations on boll damage indicated that treatment might be necessary if >20-25% reveal internal symptoms of feeding injury during mid- to late season. More detailed damage thresholds should be developed to complement an approach based on population monitoring. This study validated current recommendations for management of pentatomids in cotton, demonstrated the necessity of threshold use for stink bugs in transgenic cultivars expressing endotoxin from B. thuringiensis, and provided insight into further development of management options for pentatomids in the crop.     

Influence of Nezara viridula feeding on cotton yield, fiber quality, and seed germination

The influence of southern green stink bug, Nezara viridula (L.), adults (males and females) and fourth to fifth instars on cotton, Gossypium hirsutum L., boll abscission, seedcotton yield, fiber quality, and seed viability was evaluated in field studies conducted during 2004 and 2005. Cotton bolls representing several age classes ranging from 0-600 heat units were individually infested with a specific gender or life stage of southern green stink bug. Adults and nymphs induced abscission of bolls that accumulated 0-280 heat units after anthesis. Seedcotton yield was significantly lower in bolls infested with adults (males and females) and late instars through approximately 500 heat units after anthesis. Southern green stink bug feeding on bolls significantly affected the physical fiber properties of micronaire (measure of fiber fineness or maturity), strength, uniformity, and fiber length. Discolored cotton lint in the stink bug-infested bolls was more common than in noninfested bolls. Seed germination and development of normal seedlings for seed harvested from stink bug-infested bolls that accumulated < or =500 heat units beyond anthesis were significantly lower compared with noninfested bolls. No significant differences in boll abscission, yield, fiber quality, and seed germination were detected between southern green stink bug males and females or between adults and fourth to fifth instars.     

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.     

Composition and abundance of stink bugs (Heteroptera: Pentatomidae) in corn

The species composition and abundance of stink bugs (Heteroptera: Pentatomidae) in corn, Zea mays L., was determined in this on-farm study in Georgia. Seven species of phytophagous stink bugs were found on corn with the predominant species being Nezara viridula (L.) and Euschistus servus (Say). All developmental stages of these two pests were found, indicating they were developing on the corn crop. The remaining five species, Oebalus pugnax pugnax (F.), Euschistus quadrator (Rolston), Euschistus tristigmus (Say), Euschistus ictericus (L.), and Acrosternum hilare (Say), were found in relatively low numbers. Adult N. viridula were parasitized by the tachinid parasitoid Trichopoda pennipes (F.). There was a pronounced edge effect in distribution of stink bugs in corn. Population dynamics of N. viridula and E. servus were different on early and late-planted corn. Oviposition by females of both stink bug species occurred in mid-to-late-May and again mid-to-late-June in corn, regardless of planting date. In early planted fields, if stink bug females oviposited on corn in mid-July, the resulting nymphs did not survive to the adult stage in corn because ears were close to physiological maturity and leaves were senescing. Density of stink bug adults in early planted corn was relatively low throughout the growing season. In late-planted corn, females of both stink bug species consistently laid eggs in mid-to-late-July on corn with developing ears. This habitat favored continued nymph development, and the resulting adult population reached high levels. These results indicate that corn management practices play a key role in the ecology of stink bugs in corn agroecosystems and provide information for designing management strategies to suppress stink bugs in farmscapes with corn.     

Comparative assessment of feeding damage by pod-sucking bugs (Heteroptera: Coreoidea) associated with cowpea, Vigna unguiculata ssp. unguiculata in Nigeria

Feeding trials were conducted on three (young, mid-fill and mature) developmental stages of cowpea Vigna unguiculata ssp. unguiculata pods in the screenhouse using fourth instar nymphs and adults of Anoplocnemis curvipes (Fabricius), Riptortus dentipes (Fabricius), Mirperus jaculus (Thunberg), Clavigralla tomentosicollis St?l and C. shadabi Dolling. Anoplocnemis curvipes was observed to be the most damaging coreoid species causing a yield reduction of 26.4-51.7% followed by R. dentipes (24.4-29.4%), M. jaculus (21.9-26.9%), C. tomentosicollis (17.9-22.4%) and C. shadabi (15.9-20.4%). The fourth instar nymphs of each pod-sucking bug species caused a significantly higher cowpea yield reduction than their respective adults. Similarly, infestation on young pods compared to mid-fill and mature stages resulted in significantly higher yield reduction. The results suggest that infestation levels of two fourth instar nymphs of A. curvipes or three fourth instar nymphs of the other four pod-sucking bug species per young pod should be adequate for screening of cowpea varieties for resistance to the coreoid bugs.     

Damage by <Emphasis Type="Italic">Tibraca limbativentris</Emphasis> Stål (Pentatomidae) to Upland Rice Cultivated in Amazon Rainforest Region (Brazil) at Different Growth Stages

In this study, we evaluated the damage caused in the field by Tibraca limbativentris Stål adults at different levels of infestation (0, 1, 2, and 4 stink bugs) per three rice plants during three growth stages (V8, V13, and R4 stages) of upland rice cultivated in southwestern of Pará State, Amazon Rainforest region, Brazil. Heading time (panicle exertion) was affected by T. limbativentris infestations mainly in the vegetative stage and the whiteheads percentage in treatments ranged from 18.2 to 38%. The dead hearts percentages varied between 0 and 21.5%, and the mean number of primary branches (ramifications) ranged from 5.9?±?0.4 to 12.3?±?0.2. The number of empty spikelets was only affected in infestations with four insects/three plants, while the quantity of filled grains per panicle was affected only when infestations occurred during the vegetative stage. The total number of spikelets (filled?+?empty) per panicle decreased significantly in all phenological stages, and the percentage of damage ranged from 17 to 44% among treatments. Based on the proportion of damage observed, we suggest doubling the number of insects presently used as action threshold to 2 and 4 stink bugs per 15 stalks sampled for the vegetative stage, and of 1 or 2 stink bugs per 15 stalks sampled at the beginning of reproductive stage (R3/R4). Also, the field should be monitored during the entire vegetative stage, since most damage was observed in this phenological stage.     

Soybean seed damage by different species of stink bugs

Abstract

• 1 Damage caused by the three main species of stink bugs occurring on soybean Nezara viridula (Linnaeus), Piezodorus guildinii (Westwood) and Euschistus heros (Fabricius) was compared in field cages and in greenhouses. Infestation levels of 4 stink bugs/m row of plants (field cages) and 2 stink bug/plant (greenhouse) for 15 days during the pod filling stage are reported. At harvest, the yield and seed quality were evaluated.
• 2 In the field, there was no difference in yield between infested and insect‐free plants, but damage to seed quality varied with stink bug species. Plants damaged by P. guildinii had the lowest quality seeds. From 50 g seed samples harvested in the field, the mean weight of seeds classified as ‘good’ was 37.3 g in plants infested with P. guildinii, compared to 41.8, 44.2 and 46.6 g in plants infested with E. heros, N. viridula and the control, respectively.
• 3 Plants infested with P. guildinii showed the highest number of seeds damaged by stink bugs, whereas those infested with E. heros showed the lowest damage.
• 4 Plants infested with P. guildinii had 18.5% damaged seeds, higher than the 3.6% and 3.4% damaged seeds from plants infested with the two other species and 0.1% in control plants. The percentage of non‐viable seeds due to stink bug damage was 5.7% for P. guildinii but lower for the other two species.

     

Assessing stink bug resistance in soybean breeding lines containing genes from germplasm IAC-100

Sixty-five soybean, Glycine max (L.) Merr., breeding lines containing the stink bug resistant 'IAC-100' in their pedigrees were evaluated for their resistance to stink bug, primarily southern green stink bug, Nezara viridula L., feeding in replicated field trials from 2001 to 2005. Plots were sampled throughout the season for stink bug abundance, and, at harvest, seed samples were rated for stink bug-induced kernel damage. Individual seeds were categorized as having none, light, moderate, or heavy damage plus 100-seed wt and plot yields were determined. Both ground cloth and sweep net sampling procedures were used to compare stink bug densities between the soybean entries. Stink bug densities varied between years; however, in the years when populations exceeded four per row-meter or six per 25 sweeps, there were more damaged soybean seeds (>25%) in the entries with higher stink bug numbers. During the first 2 yr of evaluations, the mean stink bug-damaged soybean seeds ranged from 10.0 to 38.2%. From these differential responses, 28 entries were selected for continued study in 2003-2004. In 2003, stink bug-damaged soybean seeds were low, with damage ranging from 2.9 to 18.2%. In 2004, stink bug damage ranged from 8.8 to 53.2%. From these 28 lines, 12 entries were selected for an advanced field screening trial in 2005, including the IAC-100 and 'Hutcheson'. Damaged soybean seeds ranged from 18.5 to 54.1% among these 12 entries in 2005, under heavy stink bug pressure. From these evaluations, four breeding lines with either Hutcheson X IAC-100 or IAC-100 x 'V71-370' in their genealogy were identified as possible breeding material for future soybean stink bug resistance cultivar development.     

Prevalence of a trypanosomatid in the southern green stink bug, Nezara viridula

The southern green stink bug, Nezara viridula (L.), and certain of its host plants were examined to determine the prevalence and biological characteristics of an intestinal trypanosomatid. Promastigotes with short (< or = 17.5 microm excluding flagellum) and long forms (> or = 25.0 microm) usually infected < 50% of the bugs before August and > 50% (maximum 95%) during August-October, but prevalence was not host-density dependent. The flagellate was detected in adults and in all nymphal instars, at all sampling sites where at least 10 bugs were captured, and in bugs from all host plants sampled (soybean, red clover, vetch). Of bugs with flagellates, 27% were heavily infected (> 20 flagellates per 160X microscope field). Weights of infected and uninfected adults did not differ. Live flagellates were detected in bug feces and in one stem of red clover. When bugs were fed soybean pods, tomatoes, or snap beans in the laboratory, only once were flagellates detected in plant tissue (snap beans). The flagellate was cultured in modified Medium 199. This flagellate is prevalent in N. viridula populations in Louisiana and apparently does not cause significant pathological effects in N. viridula or its host plants, including soybean.     

Attractiveness of conspecific stink bugs to adult stink bug-baited traps in soybean fields

The attractiveness of live adult stink bugs used as baits in traps in soybean fields, Milyang, Korea, to conspecific stink bugs was evaluated. Both sexes of bean bug, Riptortus pedestris Fabricius (Hemiptera: Alididae), and one-banded stink bug, Piezodorus hybneri Gmelin (Hemiptera: Pentatomidae), were attracted to conspecific male adults-baited traps. Likewise, both sexes of brown-marmorated stink bug, Halyomorpha halys Stål (Hemiptera: Pentatomidae), and sole bug, Dolycoris baccarum L. (Hemiptera: Pentatomidae), were attracted to traps baited with conspecific male stink bugs. However, in Nezara antennata Scott (Hemiptera: Pentatomidae), both male and female used as baits in traps were attractive to conspecific adults. Accordingly, these results suggest that the only male adults of H. halys and D. baccarum and both sexes of N. antennata are attractive to conspecific male stink bugs.     

Development and seed number in indeterminate soybean as affected by timing and duration of exposure to long photoperiods after flowering

BACKGROUND AND AIMS: Long photoperiods from flowering to maturity have been found to delay reproductive development in soybean (Glycine max) and to increase the number of seeds per unit land area. This study was aimed to evaluate whether sensitivity to photoperiod after flowering (a) is quantitatively related to the length of exposure to long days and (b) persists throughout the whole pod-setting period. It was also evaluated whether seed number was related to changes in the duration of post-flowering phenophases. METHODS: Two field experiments were conducted with an indeterminate cultivar of soybean of maturity group V. In expt 1, photoperiods 2 h longer than natural daylength were applied during different numbers of days from the beginning pod stage (R3) onwards, while in expt 2 these photoperiod extensions were imposed during 9 consecutive days starting at different times between R3 and R6 (full seed) stages. KEY RESULTS: There was a quantitative response of development to the number of cycles with a long photoperiod. The exposure to long photoperiods from R3 to R5 (beginning of seed growth) increased the duration of R3-R6 regardless of the timing of exposure. The stages of development comprised in the R3-R6 phase were delayed by current as well as by previous exposure to long days. A positive relationship was found between seed number and the duration of R3-R6, irrespective of the timing and length of exposure to the long photoperiod. CONCLUSIONS: Sensitivity to photoperiod remained high during the reproductive period and was highly and positively coupled with the processes of generation of yield.     

Attachment ability of the southern green stink bug, <Emphasis Type="Italic">Nezara viridula</Emphasis> (L.), on plant surfaces

Traction forces of male and female southern green stink bugs Nezara viridula (L.) were measured on adaxial leaves of green beans (Phaseolus vulgaris L. cv. ‘Nagauzura’), peas (Pisum sativum L. cv. ‘Hakuryu’), rice (Oryza sativa L. cv. ‘Mac hin sung’), and soybean (Glycine max [L.] Merr. cv. ‘Enrei’), as well as on glass as a control surface. Bugs attached well on three-dimensionally structured surfaces covered with anti-adhesive epicuticular wax crystals. Strongest pulls were generated on adaxial green bean leaves, corresponding to safety factors (traction force/body weight) of 11.1 and 11.6 in males and females, respectively. These values were slightly lower on soybean (males: 7.4, females: 8.0) and rice leaves (males: 8.9, females: 10.6). Trichomes and papillae are assumed to promote pentatomid bug’s attachment. On tabular, wax-covered pea leaves, safety factors decreased significantly to 1.7 and 1.6, in males and females, respectively. Differently, on non-structured glass, safety factors resembled those on rice and soybean leaves (males: 9.6, females: 8.0). No statistical differences in traction force and safety factor between sexes were detected on any substrate. Surface wettability did not significantly affect the results. Both robust claws and tough adhesive pads enable the N. viridula bugs to grasp and adhere to a wide range of various plant substrates, including such ones covered with anti-adhesive wax crystals. However, tabular, pruinose pea leaves were detected to prevent the foothold of the southern green stink bugs, although pea is known as one of their common host plants.