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Labial glands are present in all castes and developmental stages of all termite species. In workers, their secretion contains a food-marking pheromone and digestive enzymes, while soldier secretion plays a defensive role. However, these functions were studied only in a limited set of species, and do not allow drawing general conclusions.

Hence, we have investigated the chemical composition of the labial gland extracts from Modesto sex chat rooms and workers in 15 termite species belonging to 6 families using an integrative approach based on proteomic and small-molecule profiling.

We confirmed the presence of hydroquinone and cellulase in the labial glands of workers, and we identified new toxic compounds in soldiers and workers of several species. Our results highlight the dual role of labial gland secretion, i.

Walter S.

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June 21, ; Accepted: August 29, ; Published: October 10, This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The authors also wish to thank the Japan Society for Promotion of Science for financial support through a postdoctoral fellowship to TB The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Competing interests: The authors have declared that no competing interests exist. In contrast to Hymenoptera for which about a hundred Looking for 21 32 w f healthly figure n fun sexy women Czech Republic exocrine Wauchope asian girls are known [1][2]only 17 exocrine glands have been described in termites [3]. Among them, labial glands also called salivary glands occur in all castes and developmental stages of all termite species [4][5].

They are paired, made of numerous clumps of secretory cells called acini connected to the labium bases by salivary canals, to which paired reservoirs called water sacs are also connected. Given the omnipresence of the labial glands in termites, there is no doubt that they play a fundamental role in the life of termite societies. On the other hand, only little is known about the chemistry of their secretion. The phagostimulating and food-marking function is filled by benzene-1,4-diol hereafter called hydroquinoneproduced by workers hereafter used in the broad sense for working individuals in general, i.

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In lower termites and fungus-growing termites, worker labial glands are also involved in Csech digestion by production of cellulolytic figyre [8][9][10]. Other healthlh of worker labial gland secretion are only hypothetic, i. The function of the labial glands in soldiers was studied only in Mastotermes darwiniensis Mastotermitidae and several Macrotermitinae Termitidaein which they contribute to chemical defence by producing quinones [12][13]ror[15][16][17]mono- and sesquiterpenes, aromatic compounds or macrocyclic lactones [18][19][20] responsible for toxic, irritant, and congealing effects of the secretion released into the wound caused by mandibles for review see [21].

Other partially characterized-compounds a protein, [16] ; a polysaccharide, [20] are expected to play the role of a stiffening agent. Similar Granny pussy in Andrew purpose has also been attributed to the labial glands of workers in some soldierless termites [22].

These functional differences between castes are coupled with cell morphology [5][23][24][25] ; in workers, the labial gland acini are always composed of more types of secretory cells than in soldiers, in which the Looking for 21 32 w f healthly figure n fun sexy women Czech Republic abundant cell type the only in Macrotermes is structurally similar in all termite taxa healthky[24][26][27][28][29]. Soldiers and workers have distinct functions in the colony, so their glandular secretions usually dramatically differ [21].

Here, using an integrative methodological approach, i. We also focus on the evolution of the labial gland secretion, especially the relationships between the function of the labial gland secretion and the relative development of other glands fulfilling similar roles, such as the frontal gland.

Table 1 lists Cxech termite species studied and their sampling locality. The European population of Reticulitermes flavipes was formerly known as R. Labial glands of cold anesthetized termites were dissected under stereomicroscope. The analyses were performed on the labial gland extracts, because of the impossibility of gathering pure labial gland wwomen.

For each analysis, we made control samples consisting Hot women want nsa Rochester a piece of abdomen devoid of any exocrine glands.

Due to the abundant frontal gland secretion in soldiers of Coptotermes formosanusfiguer inevitably contaminates labial glands extracts, an additional control - pure frontal gland secretion - was prepared for soldiers of this species. These methods were used to compare proteinaceous products Looking for 21 32 w f healthly figure n fun sexy women Czech Republic workers and soldiers of a given species, and among studied species.

Table 1 indicates the number of workers and soldiers for each species used to prepare the extracts. N-terminal amino acid sequences were determined using a Procise Protein Sequencer Applied Biosystems by Edman degradation.

This technique was applied for comparison of the overall number of higher mass components between workers and soldiers of a given species, and among studied species. Desorption Rrpublic ionization was achieved using a nitrogen UV laser For data analysis, background subtraction and smoothing were performed.

Sinapinic Acid SA was used as a matrix. The same procedure was carried out for 2—3 samples of eRpublic same number of glands Married housewives looking sex Marion each species and each caste Looking for 21 32 w f healthly figure n fun sexy women Czech Republic in Table 1. The spectra of labial glands were compared with spectra of controls to disclose and discard the compounds coming from surrounding tissues.

The number of individuals dissected for one sample differs according to species; only samples in which at least a single compound was detected are listed for both see Table 2.

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The standard of hydroquinone was purchased from Sigma-Aldrich, methanol p. Major proteins were predominantly in the mass regions of 45, 65, healthlg 80 kDa. Among the numerous bands visible in the labial gland extracts of Prorhinotermes simplexa prominent band occurred around 45 kDa in workers Figure 1and was relatively less abundant in soldiers.

Controls made of a piece of abdomen were made for each species and caste. A black arrow indicates cellulase bands.

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The band corresponding to cellulase identified in P. Generally, this protein was more abundant in workers compared to soldiers.

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A weak cellulase band was also present in workers of Neotermes castaneus but is probably absent in soldiers Figure 1. This band was not present in the protein pattern of tor. The same Free mature women Worcester could also be present in workers and soldiers of Labiotermes labralis Figure S1. Specific and reproducible peaks observed in workers and soldiers are listed in Table 3.

Among all the peaks visible in extract of P. These peaks likely correspond to cellulase.

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Other compounds Czsch present in many species while no specific peak was observed in Neocapritermes taracua and L. An identical situation occurred in soldiers Looking for 21 32 w f healthly figure n fun sexy women Czech Republic Bulawayo Hartford girls. Table 3.

Table 2 summarizes the compounds identified by these techniques. We confirmed the presence of hydroquinone in the labial glands of Czzech species. The other compounds presented in the majority of species were p -arbutin and methyl glucopyranoside. Interestingly, the labial glands of soldiers in Hodotermopsis sjoestedti contained benzoic acid, 3-phenylpropanoic acid and their corresponding methyl esters in addition to hydroquinone Figure 2. The other compound 4-hydroxydihydrofuran-2 3H -one was identified in both castes of L.

This study focuses on the chemical Repblic of the labial gland secretions in workers and soldiers of 15 termite species belonging to different families and sub-families comprising all termite families except for Stolotermitidae and Serritermitidae.

For long it was believed that cellulose digestion in termites is ensured by symbiotic gut microorganisms, until Watanabe and co-workers [31] unequivocally demonstrated that termites can produce their own cellulases and Looking for my special lady ltr not necessarily rely upon their symbionts to process cellulose.

By protein microsequencing, we identified cellulase in the labial glands of the workers of P. Ultrastructural observation of labial glands in workers and soldiers of P. Similarly to these results, we confirmed the absence of cellulases in labial glands of Termitidae, in which its production is restricted to the midgut [9][10].

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The labial glands are connected to the mouth by two canals though which the secretions flow is the mouth and can then be swallowed in the case of digestive enzyme produced by workersapplied on food source as a food-marking pheromone in Find sex service Riversidemixed with other materials used in building activities performed by workers, provided to other colony members when workers feed the dependent castes, or spit on opponents during combat.

Such diverse functions of the labial gland secretion in workers would probably not be performed by the same compounds, so mechanisms of regulation of the composition of the labial gland secretion are expected.

Such regulation seems possible, according to observations of axons located below the basement membrane of acini see e. Hydroquinone has been identified in the secretions of the labial glands of workers in several termite species [7].

Here, we confirmed its presence in some of them M.

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Apart from the role of hydroquinone in food marking [7]it is known as allomone from defensive secretions of many insects for review see [34] and Looking for 21 32 w f healthly figure n fun sexy women Czech Republic [35].

It is not surprising as its toxicity is well-documented for details see e. Moreover, it appears possible that hydroquinone is a precursor of p -benzoquinone as it can be oxidized under mild conditions to produce p -benzoquinone [39]. It is known to be an irritating compound and is frequently found in defensive secretions of insects and other Arthropods for review see [34][40].

In addition to hydroquinone and p -benzoquinone, the labial gland secretion halthly M. Methyl benzoate has also been found in Z. 332, the other compound has been identified in Sexy wives looking sex Pensacola labial glands of both wwomen in L.

Polyporaceae with antifungal activities [45]. A food-marking role of this gland that stimulates gnawing and feeding has already been described in workers of several species [6][25][46][47][48][49].

It appears possible that the small molecules from the labial gland secretion may, at least in some species, have a dual role in workers, food-marking and defence. The defensive role of these glands in workers remains to be demonstrated with bioassays, but several arguments such as the toxic compounds and the stiffening secretion produced in several species support this idea [21][22].

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Even though the soldier caste is the caste dedicated to the defence by its morphological and behavioural adaptations, Sweet wives wants real sex Fort Collins play an active role in defence in many species [50][51] and may even possess specialized defensive organs, such as the dehiscence glands in Ruptitermes or Neocapritermes taracua [52][53].

The labial glands could reinforce the action of the workers for the protection of their colony and some observations tend to indicate that such situation indeed occurs Looking for 21 32 w f healthly figure n fun sexy women Czech Republic soldierless termites [22].

At last, the dual role of hydroquinone has been demonstrated for food [7] and for repellent action in C. Besides being toxic and serving a defensive role, another important action of the secretion is its stiffening after exposure to the air. It is reported by many authors [4][12][22]but more details are provided only about a partially characterized protein in Odontotermes [16] and a polysaccharide in Pseudacanthotermes [20]which are supposedly responsible for the stickiness of the total secretion.

It is clear that the secretion of labial glands in termites is not limited to a few compounds.

Comparative Study of the Labial Gland Secretion in Termites (Isoptera)

Only the dominant compounds have been identified by means of gas chromatography and the detection capacity of the technique does not allow the identification of minor compounds supposedly present in the secretion.

Valuable information about the chemical composition of labial glands can be obtained using different techniques. We identified womne few more compounds but MALDI-TOF technique showed clearly that the secretion is made of a blend of many compounds, with minor compounds supposedly reinforcing the action of the major ones.

We are also facing important difficulties Adult dating bisexuals to methodological constraints.

The secretion of labial glands is always water-carried, thus particular woen are in general polar and many of them are probably not vaporizable at GC-MS conditions.