Photoperiod-dependent release of suppression pheromone in the male lobster cockroach Nauphoeta cinerea.

The complex agonistic repertoire between male lobster cockroaches (Nauphoeta cinerea) makes this species an excellent model for aggression studies. During the establishment of dominance hierarchies, 3-hydroxy-2-butanone (3H-2B) functions as a suppression pheromone, keeping the rivals in a submissive state. In the present study, we evaluated the release of 3H-2B by dominant individuals across four different time phases within the 24-h photoperiod, i.e., early scotophase (ES), late scotophase (LS), early photophase (EP), and late photophase (LP). For each time phase, we collected volatile pheromones during a 60-min first-encounter fight to measure the level of released 3H-2B. Subsequently, the amount of 3H-2B remaining in the sternal glands of dominant and subordinate individuals was measured and compared to socially naïve male controls. Release of 3H-2B was relatively high during ES or LP first-encounter fights, compared to LS or EP encounters. The attack duration and aggressive posture intensity in dominant males were positively correlated with the amount of 3H-2B release in all four phases. A similar statistical distribution was found between the amount of 3H-2B released by dominant males and the amount of 3H-2B in the sternal glands of naïve male sternal during LS, EP, and LP. However, during ES, the statistical distribution of 3H-2B released by the dominant was significantly greater than the distribution of 3H-2B content in socially naïve male sternal glands. The observed phase-dependence of 3H-2B release might be due to variations in 3H-2B biosynthesis or the scotophase-specific behavior of naïve males, wherein an aggressive posture is spontaneously adopted with concomitant 3H-2B release.

Interspecific competition and territory defense mechanisms of Coptotermes formosanus and Coptotermes gestroi (Isoptera: Rhinotermitidae).

Coptotermes formosanus Shiraki and C. gestroi (Wasmann) are the most widely distributed species of the genus and occur sympatrically in the subtropics. Results of two bioassays in the current study showed that C. gestroi was more aggressive than C. formosanus. In the petri-dish bioassays, C. gestroi won most of the agonistic encounters over C. formosanus. In the two-dimensional foraging arena bioassays, over 73% tunnel interceptions observed in the 18 replications were caused by progressing tunnels of C. gestroi encountering the tunnels of C. formosanus. Tunnel interception of the two species resulted in minor agonistic interactions. Both species quickly buried the connected tunnel at multiple locations. Termite cadavers resulting from agonistic behavior appeared to have induced sand deposition that resulted in tunnel blockages and deterred reopening of these blockages. Sealing individual tunnels in response to encounters with other species acts to prevent further agonism and mortality, and on a broad scale, the aggregate of such blocked tunnels may come to define the borders between adjacent colonies.

Tunneling response of termites to a pre-formed tunnel.

Subterranean termites move from place to place while foraging by tunneling through soil. During a period of foraging, they are likely to encounter a number of pre-formed tunnels created by, for example, tree roots or the breaking up of a zone of hard or compacted soil. We systematically observed the behavioral response of tunneling termites to such pre-formed, artificially constructed tunnels at widths, W, of 0.5, 1.0, 1.5, and 2.0mm, which mimicked pre-formed tunnels in the field. The two tunnels intersected at an angle, theta (=0 degrees , 10 degrees , 20 degrees , 30 degrees , 40 degrees , 50 degrees , and 60 degrees ) formed between the advance direction of a termite tunnel and the perpendicular direction of a pre-formed tunnel. For W=Wc (=0.5mm) and theta

Juvenile hormone levels are increased in winners of cockroach fights.

In lobster cockroach (Nauphoeta cinerea) adult males, concomitant expression of attack behavior and an increase in juvenile hormone (JH) III titer can be induced by contact with an isolated antenna [Chou et al., 2007. Antenna contact and agonism in the male lobster cockroach, Nauphoeta cinerea. Horm. Behav. 52, 252-260]. In the present study, socially naïve N. cinerea males that were either aggressive posture-adopting (i.e., "ready-for-fight") or not (i.e., "non-ready-for-fight") were paired to ask if status was determined by JH III levels before the encounter and if JH III levels were altered in dominants and subordinates after the encounter. The results showed that, although in the non-aggressive posture-adopting male pairs, the one with higher JH titers before the encounter was more likely to become the dominant, this was not the case in pairs formed between aggressive posture-adopting males or between non-aggressive posture-adopting and aggressive posture-adopting males. In all types of male pairs combined, JH III levels in the dominant were significantly increased after the encounter compared with before the encounter and were significantly higher than those in the subordinates, suggesting that the JH III increase in the dominants may serve to sustain aggression. JH III application before rank formation had a significant effect on establishment of dominant status in non-aggressive posture-adopting, but not aggressive posture-adopting, males. After rank formation, JH III application to subordinates had no effect on rank switch. These results indicate that the relationship between JH and aggression in this cockroach species is broadly consistent with the vertebrate challenge hypothesis, which predicts that testosterone levels increase in response to social stimuli to modulate aggression.

Pheromone, juvenile hormone, and social status in the male lobster cockroach Nauphoeta cinerea.

In this study, the major pheromone component, 3-hydroxy-2-butanone (3H-2B), released by dominants was measured during early scotophase. Both the JH III titer in the hemolymph and the 3H-2B content of the sternal glands of the dominants and subordinates were then measured during late scotophase and late photophase. These investigations were performed on encounter days 1, 2, 3, 5, 7, 9, 12, and 20. The results showed that, for non-aggressive posture (AP)-adopting socially naïve males (SNMs), both the 3H-2B release and the hemolymph JH III titer were maintained at a low level. Once a fight occurred, 3H-2B release was raised significantly in the AP-adopting dominants, but not in non-AP-adopting subordinates, and remained raised throughout the entire experimental period. At 30 min after the first encounter, the hemolymph JH III titer was significantly increased in dominants, but not in subordinates. A significantly higher hemolymph JH III titer was observed in dominants during late scotophase on days 3, 5, 12, and 20 and during late photophase on days 3, 5, and 20. After fighting, the sternal gland 3H-2B content of the dominants or subordinates was significantly lower than in SNMs. In dominants, the sternal gland 3H-2B content during late scotophase was significantly lower than that during late photophase in the first 9 domination days, while, in the subordinates, the 3H-2B content during late scotophase was either similar to, or significantly higher than, that in late photophase. In the dominants, 3H-2B release and JH III titer were positively correlated. In rank switchers, the switched social status was positively correlated with both 3H-2B release and JH III titer. Comparison of 3H-2B release and JH III titer in 1-time, 3-time, or 5-time dominants showed that, although winning significantly increased both 3H-2B release and JH III titer, there is no significant difference in 3H-2B release between 3- and 5-time winners, while the JH III titer was most significantly increased in the 3-time winners. The possible relationship between pheromone release, JH III titer, and social status is discussed.

Surface irregularity induced-tunneling behavior of the formosan subterranean termite.

Subterranean termites forage from place to place by tunneling through soil. In order to examine termite-tunneling responses to external factors, we designed a square arena that contains five introduction chambers connected with narrow paths, which was filled with sand. Triangle-shaped indentation with width W and height H was provided as surface irregularity on the sand facing the introduction chambers by using templates. After termites were introduced into the chambers, we measured elapsed time, tau, for a tunnel to reach the point at 3 mm away from the apex of the irregularity. We found that for W=0 mm (the absence of irregularity), termites (Coptotermes formosanus Shiraki) did not tunnel for approximately 7h, whereas, for W=1, 2, 3, and 4 mm, they exhibited the tunneling behavior within 20 min. The result indicated that the presence of surface irregularity is essential to induce termite tunneling. In addition, we found that W was correlated with tau, whereas H did not influence tau. This was briefly discussed in the context of individual movement behavior.

Antenna contact and agonism in the male lobster cockroach, Nauphoeta cinerea.

On any given day, about 35% of 80- to 85-day-old socially naïve male (SNM) lobster cockroaches (Nauphoeta cinerea) spontaneously adopted an aggressive posture (AP) without encountering another male [spontaneous AP (SAP)]. Although SAP SNMs showed significantly higher release of the pheromone 3-hydroxy-2-butanone (3H-2B) than non-SAP SNMs, there was no significant difference in hemolymph juvenile hormone (JH) III titer. When different body parts were tested for induction of the attack behavior, the antenna was found to be the most effective. After 1 min of contact with an antenna from another SAP SNM, attack behavior was induced in 100% of SAP and 76.2% of non-SAP SNMs, and the JH III titer was significantly increased in all responders. Among the non-SAP SNMs, the JH III titer before antenna contact was significantly lower in the non-responders than in the responders, and, although the JH III increase induced by 1 min antenna contact was similar between responders and non-responders, the final JH III titer of the non-responders was significantly lower. A similar attack response, JH III titer change, and 3H-2B release were seen when the individual's own antenna was used. After 5 min of contact with an antenna from another SAP SNM, attack behavior was induced in 100% of SAP and 82% of non-SAP SNMs; in the former, 3H-2B release was similar before and after antenna contact, but the JH III titer was significantly increased after antenna contact, while, in the latter, both 3H-2B release and JH III titer were significantly increased after antenna contact. Among the non-SAP SNMs, JH III titer in the non-responders was not elevated after 5 min antenna contact, and was significantly lower than that in the responders. A pentane-washed antenna did not induce attack behavior or increase the hemolymph JH III titer, and a pentane-washed antenna coated with 3H-2B also failed to induce attack behavior. These results indicate that N. cinerea male-male agonistic interactions, to which the vertebrate challenge hypothesis can be applied, are due to contact pheromone on the antenna, resulting in the concomitant expression of attack behavior and an increase in 3H-2B release and JH III titer.

Strategic 3-hydroxy-2-butanone release in the dominant male lobster cockroach, Nauphoeta cinerea.

In the lobster cockroach Nauphoete cinerea, the dominant-subordinate hierarchy formed via the agonistic interactions is unstable, and changes in rank order are common. Our previous results showed that in the first encounter fight during initial rank formation, microgram levels of 3H-2B are released by the aggressive posture (AP)-adopting dominant male. In the present study, the pattern of daily pheromone (3H-2B) release during the domination period and on the day of rank switch, rank duration, and rank switch frequency were investigated in three-male groups and six-male groups to examine the effect of higher frequency of agonistic encounters. The results showed that, in the three-male groups (50-day observation period), daily 3H-2B release rate was not constant, but fluctuated, the average duration of dominant rank was 16.6 +/- 2.0 days, rank switch occurred in 58.8% of groups, and the frequency of rank switching (average number of rank switches/group/50 days) was 1.4 +/- 0.2. For the six-male groups (30-day observation period), the daily 3H-2B release rate also fluctuated, but the duration of dominant rank was significantly shorter at 4.2 +/- 0.6 days, rank switch occurred in 100% of groups, and the frequency of rank switching (average number of rank switches/group/30 days) was significantly higher at 6.9 +/- 0.6. The results for both sets of male groups showed that as a new rank formed (either on the first encounter day or on the day of rank switching), the dominant status was significantly associated with a higher 3H-2B release rate. In the animal kingdom, fighting usually involves communication or the exchange of signals, and the results of this study indicated that the fluctuating daily 3H-2B release rate adopted by the dominants is a kind of strategic release and the 3H-2B release rate is a signal used to determine dominance.