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To evaluate the infection risk of Anoplophora malasiaca(Thomson) (Coleoptera: Cerambycidae) in two species of Japanese pine bonsais (Japanese black pine and Japanese white pine), feeding and oviposition preferences of female A. malasiacato pine bonsai and mandarin orange were examined. A female was released into a net cage together with a pair of one pine bonsai and one mandarin orange. All mandarin oranges used in preference tests were damaged by feeding and deep biting scars from females, and larval frass ejection from the trunk was found in some mandarin oranges within 4 months of observation. However, these scars were found on only a few pine bonsais, and frass ejection was not found in any pine bonsais. No A. malasiacalarvae were detected in these pine bonsais. Therefore, frass ejection from trunks might be indicative of the presence of larvae. In field surveys at bonsai nurseries, no A. malasiacaadults were found on the pine bonsais and no pine bonsais exhibited frass ejection. Statistical analyses of these results suggested that the risk of A. malasiacainfestation in pine bonsais could be ruled out as a negligible level by confirming an absence from the trunk of frass ejection.

Anoplophora malasiaca(Thomson) (Coleoptera: Cerambycidae) is a serious pest that affects various crop trees and landscapes in Japan. We collected and analyzed volatiles from male and female A. malasiaca. 4-(n-Heptyloxy)butan-1-ol and its aldehyde, pheromone components in A. glabripennisand A. chinensis,were detected in the male volatile extracts and nonanal both in the male and female volatile extracts. Nonanal was absent from the extracts of twigs of the willow host plant. Gas chromatograph-electroantennographic responses showed that nonanal and 4-(n-heptyloxy)butan-1-ol elicited responses from both male and female antennae, but 4-(n-heptyloxy)butanal did not. Volatiles of eight artificially reared males, analyzed every 3 or 4 days for 60 days from adult emergence showed that they all produced nonanal and 4-(n-heptyloxy)butan-1-ol. The two compounds produced no short-range female attraction but in males, the short-range attraction to nonanal was dose dependent and significant in a higher dose, but did not depend on 4-(n-heptyloxy)butan-1-ol. When wounded willow twigs were added to nonanal and 4-(n-heptyloxy)butan-1-ol, the frequency of male responses was higher than in all other treatments, but the same as wounded willow twigs alone. The wounded host plant willow twigs might thus be enough for male attraction. The identified volatiles from the beetles in the present study might have a weaker function for attraction from the distance compared to their host plant volatiles.

The red-necked longicorn beetle Aromia bungii(Faldermann) (Coleoptera: Cerambycidae) is an emerging invader in Japan and has become a major pest of cherry blossom trees, peaches, prunes and Japanese apricots. The male-produced aggregation-sex pheromone, recently identified as (E)-2-cis-6,7-epoxynonenal, has two enantiomers. Here, we determined which enantiomer the male produces and tested the responses of beetle antennae to the enantiomers using electroantennography. The male produced only the (6R,7S)-enantiomer, but both enantiomers elicited responses from the antennae of female beetles. Antennae of males also responded to both enantiomers. In field trials, the (6R,7S)-enantiomer and the racemic mixture were equally attractive to both sexes of the beetle, whereas the unnatural (6S,7R)-enantiomer did not attract any beetles.

The red-necked longicorn beetle, Aromia bungii(Faldermann) (Coleoptera: Cerambycidae), which has recently invaded Japan, is a serious pest species. The larvae kill cherry and orchard Rosaceae trees, such as peach, plum, and apricot. To clarify their features for mate location, male and female flight behaviors were observed in wind tunnels with caged males and females as lures. In a small wind tunnel (50 cm in diameter, 2 m in length), both sexes showed takeoff behavior according to increasing airflow. The rate of female takeoff against the male lure, a live male in a wire-netting cage, tended to be higher than for other combinations. In a large wind tunnel (ca. 1.6 m in diameter, 4 m in length), both sexes ascended and showed momentary hovering and astatic flight. When male lure cages were placed windward in the wind tunnel, females showed “upwind flight,” while males did not. It is assumed that females were induced to fly upwind by a factor derived from live males, likely to be an airborne pheromone component.

The white grub beetle Dasylepida ishigakiensisNiijima et Kinoshita (Coleoptera: Scarabaeidae) is a serious sugarcane pest on the Miyako Islands, Okinawa, Japan. Because this beetle stays underground for most of its lifetime, mating disruption using a synthetic sex pheromone has been suggested as a promising control measure. The amount of pheromone (2-butanol) released by the females is known to decrease drastically as they repeat calling each day. In this study, we determined the response of males to different concentrations of (R)-2-butanol, both in the laboratory and in the field. Males showed typical pre-mating behaviors and they were attracted to (R)-2-butanol even at a concentration 1/100 of the amount typically emitted by a female during the 1st period of calling. We examined whether female attractiveness to males was reduced with the frequency of calling, by counting the number of males attracted to females that had called during zero to five periods beforehand. Our results indicated that a high level of female attractiveness was maintained even after three periods of calling. Based on these findings, we propose that releasing synthetic sex pheromone shortly before the start of the mating season and into the late mating season may be optimal for disrupting the mating of this beetle.

The effect of mating disruption by the ground-surface application of 2-cm-long dispensers (mini-dispensers) of 2-butanol against the white grub beetle Dasylepida ishigakiensisNiijima et Kinoshita (Coleoptera: Scarabaeidae) was examined in sugarcane fields on the Miyako Island, Okinawa, Japan. Mating rates and male catches with sex pheromone traps were reduced to a low level comparable to that obtained from a conventional method in which rope dispensers (25 m) were hung at a height of ca. 30 cm along the sugarcane ridges. Both mating rates and male catches were reduced with increasing number of treated mini-dispensers. These results suggest that by using mini-dispensers the amounts of synthetic sex pheromone and plastic resin can be reduced to 1/10–1/5 and 1/6–1/3, respectively, of their conventional application with rope-type dispensers, without impairing the efficiency of mating disruption in this beetle. Furthermore, the amount of labor required for the application of this method is expected to be greatly reduced compared to the rope-type dispenser method.

We investigated physical and chemical cues involved in male mating behavior of the white grub beetle, Dasylepida ishigakiensis (Scarabaeidae). When presented with female attractant pheromone ( R)-2-butanol lures in a flight tunnel, nearly all males exhibited orientation and touching behaviors to freshly killed males and females and to intact glass models. Males landed and bent their abdomens on male and female bodies, but not on intact glass models. When treated with one female equivalent (FE) extract, washed immature male bodies and glass models both evoked stronger male responses than untreated equivalents, with the former eliciting a greater response than the treated glass models. Male responses to target male and female bodies decreased with increased numbers of washings of target bodies with organic solvents. These results suggest that the chemical factors that elicit male abdominal bending behavior are present on the body surface in both sexes. Washed immature male bodies treated with 1 FE or one male equivalent (ME) of extract induced strong male abdominal bending behavior. Washed mature female bodies treated with 1 ME extract also evoked male responses. Extracts of both sexes included factors eliciting male abdominal bending behavior. These results suggest that both physical and chemical cues derived from conspecifics cooperate to facilitate male mating recognition in D. ishigakiensis. The mating process of this species in the field is highly synchronized. Thus, after orienting to a female-like object, the only information males require by touching is whether the sex attractant pheromone that attracted them is indeed from a conspecific. [ABSTRACT FROM AUTHOR]

The use of magnetic resonance imaging (MRI) as a tool for in vivo detection of eggs in living DasylepidaishigakiensisNiijima et Kinoshita, a major pest of sugarcane, was explored using females with an ovary at different developmental stages. MRI measurements of beetles were performed at 13 °C to avoid motion artifacts on the MR images. Spin–lattice relaxation time-weighted images allowed the observation of eggs at short acquisition times (2 min, 8 s). By comparing MR images with dissection data, criteria for determining mature eggs in MR images were a clear circular or ellipsoidal shape surrounded by a relatively bright rim and a size typically larger than 1.3 mm in the minor axis. Although small oocytes could not be detected, females with a developed or undeveloped ovary could be clearly distinguished based on MR images. The possibility of confusing the digestive tract as eggs in a female with a less developed ovary can be eliminated using a proton density weighted image.

The feasibility of mating disruption with synthetic sex pheromone for the control of the white grub beetle Dasylepida ishigakiensis(Coleoptera: Scarabaeidae) was examined by permeating sugarcane fields with a racemic mixture of 2-butanol (rac-2B) released from polyethylene-tube dispensers on Miyako Island, Japan in 2011. An application of rac-2B released from 10,000 tube dispensers into 3,200 m2sugarcane fields significantly reduced the mating rate of feral females and catches of feral males with R2B-baited traps compared with the results from untreated sugarcane fields. The larval density for the treated fields was found to be nearly zero in the following winter when the corresponding figure for untreated fields was high (1.73 and 2.33/40 × 40 cm quadrat). These results clearly show that the mating disruption technique using rac-2B could be highly promising for the control of D. ishigakiensisin sugarcane fields.

The effects of changes in host plants on the mate-searching behavior and feeding preferences of the white-spotted longicorn beetle Anoplophora malasiaca(Thomson) (Coleoptera: Cerambycidae) were examined. All individuals were raised on the same artificial diet until they became pupae. Analysis of the mate-searching behavior of the males showed that many more newly emerged males were attracted to the odor of the artificial diet than to an unbaited control. We prepared three groups of beetles and fed each group on different host plants for one week. The host plants used included the following: an artificial diet (containing Morus albaLinné), Citrus unshiuMarc. branches, and Vacciniumspp. branches. The mate-searching behavior of the males changed in relation to the plant supplied for feeding. Simultaneously, the preference among the three host plants was tested. The newly emerged males preferred the artificial diet. After a week of feeding on one of the three plants, however, the adult males selected and consumed significantly more of the plant that they had just experienced than the other plants. These results suggest that the male mate-location cue can be acquired after adult eclosion. In addition, the male beetles are capable of changing their host-plant preference. If they do so, they use different odor cues for mate location. Newly emerged A. malasiacafemales showed no preference for their first choice of food among the three host plants presented, whereas the consumption was significantly larger on C. unshiubranches. After one week of feeding on different host plants, females chose their host plant after the adult stage as well as C. unshiu, but consumed mostly C. unshiu. These results suggest that the food preferences of females are different from those of males. The behavior of females may not be affected by chemical signals from their original host-plant species (as pupae) or from the host-plant species acquired when they emerge as adults.

Abstract: Males of the white grub beetle, Dasylepida ishigakiensis Niijima et Kinoshita, transfer a large amount of a colloidal substance to females during mating. In this study we investigated the effect of the male substance on the reproductive performance of mated females and the significance of multiple mating of this beetle. Females artificially separated from the males 5 min after the start of mating produced fewer eggs than those separated after 30 min or those that were undisturbed and separated spontaneously, suggesting that the male substance is used as a nutrient for egg production by the females. When females were allowed to mate with 1–4 males, multiple mating had no clear effect on reproductive performance. The amount of male substance stored in the bursa copulatrix (BC) was not significantly increased by a second mating. The functions of multiple mating of this species may be to provide a chance for females to obtain sufficient amounts of male substance when the first male to mate has only small amounts of this substance, and to increase the genetic heterogeneity of the progeny. The presence of a serine proteinase and its possible involvement in the dynamics of the BC contents are reported.

Abstract: Volatile chemicals from Citrus and Salix host plants evoke orientation behavior in males of the species Anoplophora malasiaca (Thomson) (Coleoptera: Cerambycidae). These chemicals are emitted from wounded branches. We hypothesized that when released, these chemicals may indicate the presence of an individual to other conspecifics. Insects that originate from different host plants may use different plant chemicals from their own host to communicate with conspecifics. To further explore this theory, we investigated this communication system in a population of A. malasiaca from a third host plant, blueberry (Vaccinium spp.). Males from a blueberry host (Vaccinium population) were attracted to the odor of wounded Vaccinium branches when released near a female model in the laboratory, as has been observed in males found on Citrus and Salix host plants. The Vaccinium branch extract that was attractive to the males was separated into six fractions, of which two were active. Three active compounds were subsequently identified: β-caryophyllene and sulfur from the hexane fraction, and (E)-phytol in the weakly polar fraction. The latter two active compounds of Vaccinium branches were different from those found in Citrus and Salix.

Abstract: To investigate the seasonal prevalence of Dasylepida ishigakiensis Niijima et Kinoshita (Coleoptera: Scarabaeidae), synthetic sex pheromone traps were placed on Ishigaki Island in 2008 and 2009. One population is known to occur in the forest and fly for mating from early March to early April. The trap data revealed a second population in the farmland that flew for mating much earlier (mid-January to late February) than did the other population, indicating the presence of allochronic populations. Adults of the farmland population were caught throughout the sugarcane-grown area on the island, whereas those of the forest population appeared to be limited to the forest area. The temporal pattern of trap catches of each population was generally the same between 2008 and 2009. Relatively large trap catches were observed in the west central plane area and a small northern area of the northeastern peninsula of the island. These results suggest that, although it has not caused any damage to agricultural crops on this island, this beetle might be a potentially serious pest of sugarcane crops in the future, as reported on another island of Okinawa.

Abstract: Virgin females of the white grub beetle, Dasylepida ishigakiensis Niijima et Kinoshita, emit the sex pheromone, 2-butanol. To investigate the relationships between pheromone emission and ovarian development in this beetle, virgin adult females were kept at 18°C for various lengths of time. The sex pheromone released by individual females at first calling was collected at 23°C and the amounts determined. Females started developing visible oocytes at day 44 and emitting sex pheromone at day 50. Almost all females at day 52 or later possessed eggs, indicating that diapause had ended. A high positive correlation was found between the amounts of pheromone emitted and levels of ovarian development. A large variation of pheromone emission was encountered even among individuals of the same age. Females retained the ability to emit sex pheromone as long as they were virgin. These results may provide important information regarding the timing and duration of application of synthetic sex pheromone in an attempt to control this pest through disrupting the male-female communication.

Abstract: In 2010, abrupt outbreaks of the African armyworm, Spodoptera exempta (Walker), occured on the Tarama, Iriomote and Kikai Islands in southwestern Japan. Analysis by gas chromatography-electroantennographic detection (GC-EAD) revealed two EAG-active compounds on male antenna in crude extract of virgin females. These compounds were identified as (9Z)-9-tetradecenyl acetate (Z9-14:Ac) and (9Z,12E)-9,12-tetradecadienyl acetate (Z9E12-14:Ac) in ca. 90:10 ratio by subsequent GC-MS analyses. (11Z)-11-Hexadecenyl acetate (Z11-16:Ac), which had previously been identified as a third component in the Kenyan population, was not detected. Binary blends of Z9-14:Ac and Z9E12-14:Ac at ratios between 99:1 and 90:10 showed a potent attractiveness in the field, superior to that of virgin females and comparable to that of the three-component formulation determined in Kenya. For the population survey, a 98:2 blend was used. In Tarama, only a few moths of S. exempta were captured with a light trap during the night when more than 600 males were captured with synthetic sex pheromone; more S. exempta captures with a light trap had been reported than with sex-pheromone traps in Kenya. This indicates that the Okinawan population has different properties from the Kenyan population in pheromone composition and behavioral response to light.

Abstract: Genetic variation of the white-spotted longicorn beetle Anoplophora spp., distributed on the Japanese mainland and the Ryukyu Islands, was examined using nucleotide sequences of mitochondrial DNA. Two fragments, a 1.2-kb-long fragment containing portions of the cytochrome oxidase subunit I and II genes and a 1.4-kb-long fragment containing portions of 16S and 12S rDNAs, were sequenced. In phylogenetic analyses based on the sequences, 294 individuals collected at 75 localities were divided into two major groups (groups A and B) and then split into seven subgroups (A1–A4 and B1–B3). In group A, the closely related subgroups A1, A2, and A3 were mainly distributed on the Japanese mainland and were roughly separated among geographic areas, although the range of A3 spread significantly from Kyushu to the Central Ryukyu Islands. The sequence of A4, detected from one individual collected in eastern Honshu, was almost the same as that reported for A. chinensis (Forster) on the Chinese continent. In group B, subgroups B1 and B2 were restricted to the Central and Southern Ryukyu Islands, respectively, while B3 was distributed widely in both regions. Based on the results, we discuss the geographic origin of the haplotypes and movement of the insects between geographic areas.

Abstract A single component in extracts of virgin female Milionia basalis pryeri moths elicited responses from male moth antennae. This compound (ca. 7 ng/female) was identified as (Z,Z)-(3S,4R)-3,4-epoxynonadeca-6,9-diene by GC-MS and NMR analyses, microchemical reactions, and comparative chiral HPLC. In a field test, synthetic (Z,Z)-(3S,4R)-3,4-epoxynonadeca-6,9-diene attracted male moths.The opposite enantiomer, the racemic mixture, and virgin female moths held in small cages attracted no more moths than the solvent controls.

Two distinct electroantennographycally active (EAG-active) components, A and B, and a weakly active component C were found in a solvent extract from virgin females of the Ishigaki strain of the tussock moth, Orgyia postica(Walker). Components A, B, and C were found in the extract of the females at 4.0, 0.5, and 4.0 ng/female respectively. Components A, B, and C were identified as (6Z,9Z,11S,12S)-11,12-epoxyhenicosa-6,9-diene [(11S,12S)-1: posticlure], (6Z)-henicos-6-en-11-one (2), and (6Z,9Z)-henicosa-6,9-diene (3), respectively. Component B was absent in the extract from the Okinawa strain, in which components A and C were present at 2.0 and 1.5 ng/female respectively. (11S,12S)-1and the racemic mixture showed attractiveness for both the Okinawa and Ishigaki strains, whereas (11R,12R)-1did not. The addition of 2significantly reduced the trap catches with (11S,12S)-1on the Okinawa strain which lacked 2, while there was no significant inhibitory effect on the Ishigaki strain. The addition of 3to (11S,12S)-1did not significantly affect trap catches at Ishigaki or Okinawa. This confirmed that the attractant pheromone of O. posticaof the Ishigaki strain is also (11S,12S)-1.

Summary. Under field conditions significantly more black chafer, Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae) males landed on black and gray sources than white ones containing 10 mg of female pheromone, anthranilic acid. When a broader spectrum of colours was tested the frequency of male landing was intensively proportional to as the lightness of the colour of the lures. These findings demonstrated that mate location by H. l. loochooana males is dependent on both olfactory and visual stimuli of the source. In contrast, female aggregation was not affected by colour.

Summary. Both male and female Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae) were attracted with female-produced pheromone, anthranilic acid (2-aminobenzoic acid), in the field. Male chafers were observed to apparently directly locate cotton balls impregnated with 1 to 10 mg of pheromone. In contrast, females never directly oriented to the treated balls but landed 0.2-1.5 m away and exposed their abdominal glands in a calling posture, which occasionally resulted in aggregation of both females and males. This suggested the mating aggregation of this species could be primarily induced by pheromone released by females. A hypothesis for adaptability of female aggregation is proposed and discussed.

Summary.: Ether extract of the abdominal gland of female black chafers Holotrichia loochooana loochooana (Sawada) (Coleoptera: Scarabaeidae) induced a series of pre-mating behaviors that included short-distance orientation, mounting and abdominal bending toward females in the laboratory. When the extract was chromatographed on a silica gel column, the activity was eluted with 50% ether in hexane and ether fractions. GC-MS analyses revealed that both the active fractions contained anthranilic acid (2-aminobenzoic acid) as a major compound. When partitioned to basic, acidic and amphoteric fractions, pheromonal activity was observed with the latter two fractions and anthranilic acid was detected in these fractions by HPLC analyses. The amount of anthranilic acid in the female extract was estimated to be ca. 1.3 μg/female. Authentic anthranilic acid induced pre-mating behavior in males of at levels equal to that elicited by the compound in the extract. It also attracted male beetles to the traps in the field.

Two EAG‐active compounds were found in the solvent extract of abdominal tips of virgin females of the tussock moth Euproctis pulverea(Leech) (Lepidoptera: Lymantriidae), and identified as (Z,Z,Z)‐11,14,17‐icosatrienyl isobutyrate and (Z,Z,Z)‐11,14,17‐icosatrienyl 4‐methylvalerate at 190 and 80 ng female−1, respectively, by means of GC‐MS analyses and chemical derivatization. Esters of n‐butyric acid, n‐valeric acid, n‐hexanoic acid and a methylheptanoic acid were also found at 3, 2, 0.4 and 9 ng female−1as minor EAG‐inactive compounds. Two active compounds were also detected in the hexane extract of female anal tufts at 17 and 6 ng female−1, respectively. In Okinawa, the binary blend of the synthetic compounds attracted male moths to the sticky traps, but single compounds did not. The significance of these findings in relation to parasitism by Telenomus euproctidis(Hymenoptera: Scelionidae) is discussed.

A predatory stink bug, Eocanthecona furcellata, died after feeding on Milionia basalislarvae. The compounds toxic to E. furcellatawere isolated from the hemolymph of M. basalislarvae and identified as inumakilactone A, nagilactone C, and nagilactone C glucoside. The concentrations of inumakilactone A, nagilactone C, and nagilactone C glucoside in the hemolymph of the final instar larvae were 130, 50, and 770 μg/ml, respectively. Nagilactone C showed the highest insecticidal activity against second-instar nymphs of E. furcellata, while nagilactone C glucoside showed the lowest, one twentieth of that of nagilactone C. When mixed compounds were given at the same concentrations as those in hemolymph of M. basalis, all nymphs of E. furcellatadied with in three days. Inumakilactone A and nagilactone C were found to be in the leaves of podocarp, Podocarpus macrophyllus, the only host plant of M. basalis, at concentrations of 13 and 175 μg/g fresh weight, respectively. However, no nagilactone C glucoside was detected in the leaves of this species. These results suggested that M. basalismay transform nagilactone C to its glucoside.

In previous studies, we reported the existence of a high concentration of free glycine in the digestive juice of several Lepidoptera larvae, particularly in the digestive juice of species that feed on the privet tree, Ligustrum oblusifolium.The water extract of privet leaves showed very strong protein-denaturing activity and lysine-decreasing activity, which closely resembled activity of oxidized polyphenolics. Addition of 1% glycine to the extract could completely inhibit these activities. Free glycine may be secreted into the digestive juice by larvae as an adaptive mechanism for chemical defense against its host plants. The protein-denaturing compound in privet leaves is present in the cytosol or in the vacuoles of the leaf cells. The compound does not show protein-denaturing activity without oxidation, but when mixed with intact organelles under low osmotic conditions to give an osmotic shock, a very high protein-denaturing activity is produced. Our results suggest that the privet tree is endowed with a defense mechanism in which a stable compound in the cytosol or in the vacuoles is activated into a chemically active denaturant by an enzyme present in the organelles (including chloroplasts) after the leaves are eaten by insects and the organelles are broken by osmotic shock or by digestive mechanisms. Based upon HPLC and NMR data, we conclude that the denaturing compound is oleuropein, an o-dihydroxyphenolic compound. This compound makes up 3% of the wet weight of privet leaves. The protein-denaturing activity of purified oleuropein activated by the leaf enzyme is high enough to account for all the denaturing activity in a water extract of privet leaves. The denaturing reaction is completely inhibited by free glycine. Our results suggest that the protein-denaturing activity and lysine-decreasing activity of privet leaves are caused by oxidized polyphenolics, and that some insects secrete free glycine to counter the denaturing activity of oxidized phenolics. The chemical mechanism of counteraction by glycine is also considered. Free glycine in the midgut of insects probably protects proteins from denaturation by competing with proteins for oxidized phenolics.

Bitter gourd, Momordica charantia, was less palatable to two species of armyworms, Spodoptera lituraand Pseudaletia separata, than two other cucurbitaceous plants. A methanol extract of M. charantialeaves inhibited feeding of the armyworm larvae. The two most active fractions obtained by silica gel chromatography were purified by HPLC. Momordicine II, a triterpene monoglucoside, was identified as an antifeedant compound from the more active of these fractions. The second active fraction led to the isolation of a new triterpene diglucoside. Fresh leaves of M. charantiacontained ca. 0.3% of momordicine II. Momordicine II showed significant antifeedant effects on P. separataat concentrations of 0.02, 0.1, and 0.5% in artificial diets. Momordicine II caused a significant feeding reduction in S. lituraonly at the highest concentration (0.5%) tested. The difference in feeding response of the two armyworms to momordicine II may be related to the diversity in their host range.

(+)-Boronolide (= (1′R,2′R,3′S,6R)-5,6-dihydro-6-[1′,2′,3′-tris(acetoxy)heptyl]-2H-pyran-2-one) (1), a constituent isolated from Tetradenia fruticosaand T. barberae(Lamiaceae), was synthesized in 7 steps and in 5.2% overall yield from 2,3,4,5,6-penta-O-benzyl-d-glucose, whose four asymmetric centers were incorporated in 1.

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