Paper wasp/Common paper wasp

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Code: i4
Latin name: Polistes spp.
Source material: Venom
Family: Vespidae
Paper wasp includes 5 North American species.
 
See also European paper wasp/Mediterranean paper wasp (Polistes dominulus) i77.
 
Venom
An insect, which may cause allergy symptoms in sensitised individuals.
 

Allergen Exposure

Geographical distribution
Insects of the order Hymenoptera are mainly responsible for causing allergic reactions from stings. However, only species belonging to 3 families sting people with a high degree of frequency: Bees (Apidae), Ants (Formicidae), and species of the family Vespidae, to which belong Wasps (Polistes), Hornets (Vespa/Dolichovespula), and Yellow jackets (Vespula) (1-4).

The genera Vespula, Dolichovespula and Vespa are found all over Europe. The genus Vespula predominates over Polistes and Vespa throughout Europe, except in Mediterranean areas. Polistes is the most common genus among social Wasps and has a worldwide distribution. In Europe and North America their colonies outnumber those of all other social Wasps combined (5); although they are present in central Europe, they are not found in the UK and are rare in northern Europe (4).

P. dominulus, P. gallicus, P. biglumis and, to a lesser extent, P. ninphus are the most widely spread in Europe and are especially relevant in Mediterranean areas, while P. annularis, P. fuscatus, P. metricus, P. apachus and P. exclamans are present in the USA (5-7). There are more than 20 species of Polistes found within the United States, the greatest diversity being in the southeast (7). The American species are not present in Europe. In 1981, P. dominulus (European species) was first reported in the Boston and New York City areas and has since spread across the continent. It is now the predominant Paper wasp found there (8-10).  It has also been reported in Australia. Its invasive success is attributed to establishment of colonies earlier in the spring, reuse of abandoned nests, and a more varied diet than the native species has (11).

Recent systematic studies have confirmed that North American and European species of Polistes belong to different subgenera and are phylogenetically distant. The most common European species - P. dominulus, P. gallicus, P. nimphus and P. biglumis - belong to the subgenus Polistes sensu stricto, while the most common North American native species - P. exclamans, P. annularis, P. metricus and P. fuscatus - are included in the subgenera Aphanilopterus and Fuscopolistes (12-13).

The cross-reactivity between European and American Polistes is only partial (14). (See below)
 
Environment
Compared to other Hymenoptera, Paper wasps have a more primitive social organisation. The nests, with simple, single-layer paper cones, a maximum of about 100 cells, and no protective outer covering, are attached by a pedicle and have relatively low populations.

Fertilised queens hibernate during the winter in secluded places, but other wasps in the colony may visit the old nest on warmer days to access the honey caches there (15). One queen (or several, but in this case there will be a single dominant egg-layer) starts a nest in the spring by laying eggs, and by late July there are up to 200 workers available. The first female workers tend to be sterile and smaller, but later broods are fertile, providing for several queens. Sexual larvae are produced starting in August, and in September adults (including the only recently produced males) leave the nest and mate outside (8-11). Mated females then seek a safe place to hibernate. Polistes has a relatively simple life cycle, which is well suited to a warm climate such as in the Mediterranean (4).

Paper wasps tend to build nests on sheltered and little-used parts of buildings, placing humans in inadvertent danger of stings. Paper wasps can also be scavengers – often congregating around food factories and food shops - and this also brings them into human contact (8). They are, however, less aggressive than other members of the Vespidae. But it should be kept in mind that, whereas Bees lose their barbed stingers in a single attack and die, all Wasps, Hornets, and Yellow jackets can sting many times.
 
Unexpected exposure
See under Environment.
 
Allergens
The amount of venom released during a sting varies from species to species and even within the same species. Bee stings release an average of 50 ug (16) to 140 ug (17) of venom protein per sting; however, venom sacs may contain more than 300 ug of venom (18). Bumble bee stings release 10–31 ug of venom (16). In contrast, Vespinae, which are capable of repeated stings, generally inject less venom per sting: Vespula stings release 1.7–3.1 ug, Dolichovespula stings 2.4-5.0 ug, and Polistes stings from 4.2 to 17 ug of protein (16). One set of observations indicates that least 90% of the venom sac content is delivered within 20 seconds, and that the venom delivery is complete within 1 minute, suggesting that a stinger must be removed within a few seconds after the sting to prevent anaphylaxis in an allergic person (17).

Though not all allergens in the venom of each vespid species have been characterised, 4 groups of allergens have been identified: group 1 consists of forms of phospholipase A1, group 2 of hyaluronidase, group 4 of serine protease, and group 5 of antigen 5 (Ag5), a protein of uncertain function (19-21). The 3 most important allergens of venom from vespid Wasps are hyaluronidase, phospholipase A1 and antigen 5. They have molecular weights of about 44, 36 and 22 kDa respectively (22).

As this ImmunoCAP tests for a mixture of American Polistes species, the following allergens will be present:
  • Pol e 1, phospholipase A1, a lipase (23).
  • Pol e 2, a hyaluronidase.
  • Pol e 4, a trypsin-like serine protease (23-25).
  • Pol e 5, previously known as Ag5 or antigen 5 (23, 26-27).
     
  • Pol a 1, phospholipase A1, a lipase (28-29).
  • Pol a 2, a hyaluronidase (28).
  • Pol a 5, previously known as Ag5 or antigen 5 (26, 30-32).
     
  • Pol f 5 (33).
  • Pol m 5 (23).
The biological function of antigen 5 (Pol e 5) is not known, although it has been reported that antigen 5 from Vespa mandarinia is a neurotoxin active at the neuro-muscular junctions of a Lobster walking leg (34).
 

Potential Cross-reactivity

Studies within the Polistes genus show extensive cross-reactivity among various species (6, 35-36). However, the European species of Polistes are different from those found in America. Significantly and surprisingly, the cross-reactivity between European and American Polistes is only partial, with significant differences in specificities (5, 14, 21, 28, 37). Skin-specific IgE evaluation and direct RAST have confirmed these results (14).

Polistes is more distantly related to Vespula, Vespa, and Dolichovespula, which have a high degree of cross-reactivity, and its cross-reactivity is generally lower than within Vespinae (4, 38-40).

Vespids (Wasps, Hornets, and Yellow jackets) each have unique as well as homologous venom allergens. The homologous venom allergens have varying degrees of sequence identity, ranging from about 60% for phospholipases and antigen 5s to about 80% for hyaluronidases. Amino acid sequence similarities between Polistes and other vespid allergens are less marked (1).

In general, phospholipases of Hornets, Yellow jackets and Wasps have a 54 to 68% sequence identity, their hyaluronidases a 73 to 92% sequence identity, and their antigen 5s a 58 to 67% sequence identity (1, 41).

The cross-reactivity of the phospholipase (PLA1) venom allergen varies considerably among species. For example, PLA1s from 2 species of Yellow jackets, Vespula maculifrons and Vespula vulgaris, have 95% sequence identity with each other but only about 67 and 55% identity with White-face hornet and Paper wasp proteins, respectively (1, 42). There is also limited cross-reactivity between Polistes and the other vespid venoms, because of differences in the epitopes on the allergen molecules (43). In a study of the major allergens (phospholipase, antigen 5, hyaluronidase and protease) of Polistes gallicus (European species) with those of Polistes annularis (American species), an 85% sequence identity was demonstrated between either Ag5, which increased to 98% within the same subgenus. The authors suggested that this may be a result of the common presence or variations of specific epitopes on the Ag5 molecule (43). Still other studies have reported that there are multiple antigenic determinants on the phospholipase molecules, that individuals respond to different determinants, and that no general patterns of cross-reactivity could be observed (44).

Interestingly, the amino acid sequence of the phospholipase from Fire Ant, Sol i 1, compared with the sequences of 8 other venom phospholipases, had a higher homology with Pol d 1.01 (285 residues), as opposed to a 35% identity with Pol a 1 (209 residues) (45).

The protease allergens of Paper wasp, e.g., Pol a 4, are important. Although serine protease allergens from Honey bee, Bumble bee, and Paper wasp have been shown to have significant IgE binding activity, the structures are poorly conserved, suggesting little cross-reactivity (46).

Pol a 5, or Ag 5 (antigen 5), is a major allergen of vespid venom. A high degree of cross-reactivity of Hornet Ag 5 with Wasp or Yellow jacket Ag 5 and a low degree of cross-reactivity of Yellow jacket Ag 5 with Wasp Ag 5 was reported (26). Other studies have also reported varying degrees of cross-reactivity among homologous antigen 5 from a large number of different Yellow jackets, Hornets, and Paper wasps, with patients demonstrating varying extents of cross-reactivity to the related antigen 5s (47).

Ag 5 from Yellow jacket (Vespula vulgaris) and P. annularis (American species) have a 59% sequence identity and 69% with White-face hornet (Dolichovespula maculate), and low degrees of antigenic cross-reactivity in insect-allergic patients and in animal models (29, 32). There is a greater degree of cross-reactivity among the homologous allergens of Hornets and Yellow jackets than among the homologous allergens of Hornets or Yellow jackets and Wasps (32).

This is in the same pattern as that reported for the frequency of patients’ multiple sensitivity to these insects (48). However, a study investigating the degree of cross-reactivity between Vespula vulgaris and P. dominulus (European species) reported that in 31 of 45 patients, the double sensitisations to venoms were probably the result of cross-reactions (49).

However, apart from phospholipases (e.g., Pol e 1), antigen 5s (e.g., Pol e 5) are a major cause of the different immunological responses between American and European venoms. Although there is a 98% sequence identity between antigen 5 derived from the European species of Paper wasp P. dominulus and P. gallicus, the identity between the antigen 5 of P. gallicus and that of the American paper wasps P. annularis and P. exclamans decreases to 85%, and although this is still technically "high", the difference confirms the observed poor cross-reactivity between American ad European Paper wasps (46).

Similarly, even though a high degree of homology could be shown between 2 forms of antigen 5 of Polybia scutellaris and those of other vespids, amino acids at positions 5 and 11 in the P. scutellaris
antigens differed from the previously known sequences for antigen 5, suggesting that one or the other amino acid might be responsible for the lack of allergenicity of the P. scutellaris venom (50). This confirms the variability of the antigenicity and cross-reactivity of antigen 5. Similarly, although Fire ant (Solenopsis invicta) contains an antigen 5 venom allergen, it lacks IgE cross-reactivity (33).

A 43-45 kDa hyaluronidase protein of Vespula vulgaris and similar proteins from 5 other Vespula Wasp species have been isolated and cloned, and found to have a 59% homology to P. annularis hyaluronidase (Pol a 2) (51).

Clinical expression of cross-reactivity between Vespula and Polistes venom allergens is illustrated in a study of 28 Spanish patients from Madrid and its surroundings, who had experienced systemic reactions to vespid stings. All patients had serum-specific IgE directed at Vespula venom, and half of them had it directed at Polistes venom. Polistes-specific IgE could be inhibited with either Polistes or Vespula venom to a similar degree, whereas inhibition of Vespula-specific IgE was possible with Vespula venom, but only to a limited degree with Polistes venom. The authors concluded that in this geographic region, sensitisation to Vespula venom was more common than to Polistes venom, and that Polistes may have cross-reactivity in these patients. Therefore, an individual can be sensitive to both venoms either due to a clinical sensitisation to both venoms, or due to common antigenic determinants (52).
 

Clinical Experience

IgE-mediated reactions
Paper wasps commonly induce symptoms of food allergy in sensitised individuals (53-54). Clinical signs of Wasp stings include erythema, oedema, and pain at the sting site. Reactions may remain local, or become regional, or become generalised with systemic anaphylactic responses. Less commonly, delayed-type hypersensitivity may occur (55). Anaphylactic reactions to Hymenoptera stings are not dose-dependent or related to the number of stings (3, 14, 28). Most deaths from Hymenoptera stings are the result of immediate hypersensitivity reactions that result in anaphylaxis. However, massive envenomations can cause death in non-allergic individuals. The lethal dose has been estimated to be approximately 20 stings/kg in most mammals (2).

Systemic allergic reactions to Hymenoptera venom in general have been estimated to occur in 0.4 to 3.3% of a population. Epidemiological investigation indicates that from 15 to 25% of the general population can be sensitised to different Hymenoptera venoms, and the degree of exposure may be related to the different prevalences found in those studies (56). Older United States studies suggested that deaths from Polistes stings were more common than those from Vespula or Dolichovespula (57-58). The European experience is different, with Honey bee and Vespula deaths more common (3).

However, the prevalence of hypersensitivity to Polistes is unclear: in some studies of Hymenoptera stings, the term Wasp was used but no specific details were given about the species involved (59). Some studies grouped Polistes species together (60). This adds greatly to the confusion about the role of different vespids in sting events. In other studies, only one vespid was studied, although other species were present in the country (4, 61). Nonetheless, some guidance and information can be gleaned from a number of other published studies. Importantly, regional differences may occur due to a number of factors relating to the insects’ distribution and interaction with human activities (such as Bee keeping). Methods of detecting sensitisation also vary widely. In the central, northern and eastern parts of Europe, Vespula, Vespa and Bee stings appear to be most prevalent, whereas in the Mediterranean areas, it is Polistes and Vespula stings (4). Furthermore, since the cross-reactivity between European and American Polistes is only partial, with significant differences in specificities, studies may not be extrapolated between the pair; this has been confirmed in in vitro and in vivo studies (5, 14, 28, 37, 43). Indeed, in a study of 4 Polistes-allergic individuals in whom skin-specific IgE was negative to the American Polistes, all were positive to European Polistes venoms; 2 had  experienced severe reactions (14). The authors concluded that there is no correlation between skin test reactivity and severity of the reaction. (See European paper wasp/Mediterranean paper wasp [Polistes dominulus] i77 for an overview of European Polistes.)

Epileptic attacks have been attributed to Wasp stings. In the case of a patient who had experienced 2 epileptic attacks, both associated with anaphylaxis due to Wasp venom, and in whom serum-specific IgE against Polistes species venom was raised, it was considered likely that the attacks were due to anoxia secondary to hypotension caused by vasodilatation and cardiac involvement in anaphylaxis (62).
 
Other reactions
Electrographic changes after insect stings have been reported both with and without systemic symptoms. A patient is described who developed paroxysmal atrial fibrillation after receiving venom and pollen immunotherapy. The patient was found to be in atrial fibrillation and to have systemic symptoms immediately after administration of immunotherapy. After an injection of epinephrine, the patient converted back to normal sinus rhythm and the systemic symptoms resolved (63).

Although unusual, delayed clinical reactions to Hymenoptera stings have been reported: they can affect organs or systems generally not involved in the immediate hypersensitivity reaction. A 51-year-old man was reported as having presented with immediate, strong local and systemic allergic reactions after 2 stings of Paper wasps; the second set of reactions was triggered by a string that occurred after the third venom immunotherapy injection. He quickly improved after intravenous methylprednisolone administration, but developed acute polyradiculoneuropathy with muscle weakness, paresthesia and difficulties in standing up and walking approximately 40 hours later. Skin and specific IgE tests showed sensitisation to Paper wasp. Based on laboratory findings, the authors suggested that the mechanism may have been a delayed immunological response to Wasp antigens, followed by an allergy-triggered autoimmune reaction (64).

Compiled by Dr Harris Steinman, harris@zingsolutions.com

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As in all diagnostic testing, the diagnosis is made by the physican based on both test results and the patient history.