Biological weapons may be employed in various ways to gain a strategic ortactical advantage over the enemy, either by threats or by actual deployments. Like somechemical weapons, biological weapons may also be useful asarea denial weapons. These agents may be lethal ornon-lethal, and may be targeted against a single individual, a group of people, or even an entire population. They may be developed, acquired, stockpiled or deployed bynation states or by non-national groups. In the latter case, or if a nation-state uses itclandestinely, it may also be consideredbioterrorism.[9]
Biological warfare and chemical warfare overlap to an extent, as the use oftoxins produced by some living organisms is considered under the provisions of both the BWC and theChemical Weapons Convention. Toxins andpsychochemical weapons are often referred to asmidspectrum agents. Unlike bioweapons, these midspectrum agents do not reproduce in their host and are typically characterized by shorter incubation periods.[10]
A biological attack could conceivably result in large numbers ofcivilian casualties and cause severe disruption toeconomic and societal infrastructure.[11]
A nation or group that can pose a credible threat of mass casualty has the ability to alter the terms under which other nations or groups interact with it. When indexed to weapon mass and cost of development and storage, biological weapons possess destructive potential and loss of life far in excess of nuclear, chemical or conventional weapons. Accordingly, biological agents are potentially useful as strategic deterrents, in addition to their utility as offensive weapons on the battlefield.[12]
As a tactical weapon for military use, a significant problem with biological warfare is that it would take days to be effective, and therefore might not immediately stop an opposing force. Some biological agents (smallpox,pneumonic plague) have the capability of person-to-persontransmission viaaerosolizedrespiratory droplets. This feature can be undesirable, as the agent(s) may be transmitted by this mechanism to unintended populations, including neutral or even friendly forces. Worse still, such a weapon could "escape" the laboratory where it was developed, even if there was no intent to use it – for example by infecting a researcher who then transmits it to the outside world before realizing that they were infected. Several cases are known of researchers becoming infected and dying ofEbola,[13][14] which they had been working with in the lab (though nobody else was infected in those cases) – while there is no evidence that their work was directed towards biological warfare, it demonstrates the potential for accidental infection even of careful researchers fully aware of the dangers. While containment of biological warfare is less of a concern for certain criminal or terrorist organizations, it remains a significant concern for the military and civilian populations of virtually all nations.
Rudimentary forms of biological warfare have been practiced since antiquity.[15] The earliest documented incident of the intention to use biological weapons is recorded inHittite texts of 1500–1200 BC, in which victims of anunknown plague (possiblytularemia) were driven into enemy lands, causing an epidemic.[16] The Assyrians poisoned enemy wells with the fungusergot, though with unknown results.Scythian archers dipped their arrows and Roman soldiers their swords into excrements and cadavers – victims were commonly infected bytetanus as result.[17] In 1346, the bodies ofMongol warriors of theGolden Horde who had died ofplague were thrown over the walls of thebesieged Crimean city of Kaffa. Specialists disagree about whether this operation was responsible for the spread of theBlack Death into Europe, Near East and North Africa, resulting in the deaths of approximately 25 million Europeans.[18][19][20][21]
Biological agents were extensively used in many parts of Africa from the sixteenth century AD, most of the time in the form of poisoned arrows, or powder spread on the war front as well as poisoning of horses and water supply of the enemy forces.[22][23] InBorgu, there were specific mixtures to kill,hypnotize, make the enemy bold, and to act as an antidote against the poison of the enemy as well. The creation of biologicals was reserved for a specific and professional class of medicine-men.[23]
During theFrench and Indian War, in June 1763 a group ofNative Americans laidsiege to British-heldFort Pitt.[24] Following instructions of his superior, ColonelHenry Bouquet, the commander of Fort Pitt,Swiss-born Captain Simeon Ecuyer, ordered his men to take smallpox-infested blankets from the infirmary and give it to a Lenape delegation during the siege.[25][26][27] A reported outbreak that began the spring before left as many as one hundred Native Americans dead inOhio Country from 1763 to 1764. It is not clear whether the smallpox was a result of the Fort Pitt incident or the virus was already present among theDelaware people as outbreaks happened on their own every dozen or so years[28] and the delegates were met again later and seemingly had not contracted smallpox.[29][30][31] During theAmerican Revolutionary War,Continental Army officerGeorge Washington mentioned to theContinental Congress that he had heard a rumor from a sailor that his opponent during theSiege of Boston, GeneralWilliam Howe, had deliberately sent civilians out of the city in the hopes of spreading theongoing smallpox epidemic to American lines; Washington, remaining unconvinced, wrote that he "could hardly give credit to" the claim. Washington had already inoculated his soldiers, diminishing the effect of the epidemic.[32][33] Some historians have claimed that a detachment of theCorps of Royal Marines stationed inNew South Wales, Australia, deliberately usedsmallpox there in 1789.[34] Dr Seth Carus states: "Ultimately, we have a strong circumstantial case supporting the theory that someone deliberately introduced smallpox in the Aboriginal population."[35]
By 1900 thegerm theory and advances inbacteriology brought a new level of sophistication to the techniques for possible use ofbio-agents in war. Biological sabotage in the form ofanthrax andglanders was undertaken on behalf of theImperial German government duringWorld War I (1914–1918), with indifferent results.[36] TheGeneva Protocol of 1925 prohibited the first use of chemical and biological weapons against enemy nationals in international armed conflicts.[37]
With the onset ofWorld War II, theMinistry of Supply in theUnited Kingdom established a biological warfare program atPorton Down, headed by the microbiologistPaul Fildes. The research was championed byWinston Churchill and soontularemia,anthrax,brucellosis, andbotulism toxins had been effectively weaponized. In particular,Gruinard Island in Scotland, was contaminated with anthrax during a series of extensive tests for the next 56 years. Although the UK never offensively used the biological weapons it developed, its program was the first to successfully weaponize a variety of deadly pathogens and bring them into industrial production.[38] Other nations, notably France and Japan, had begun their own biological weapons programs.[39]
When the United States entered the war, Allied resources were pooled at the request of the British. The US then established a large research program and industrial complex atFort Detrick, Maryland, in 1942 under the direction ofGeorge W. Merck.[40] The biological and chemical weapons developed during that period were tested at theDugway Proving Grounds inUtah. Soon there were facilities for the mass production of anthrax spores,brucellosis, andbotulism toxins, although the war was over before these weapons could be of much operational use.[41]
The most notorious program of the period was run by the secretImperial Japanese ArmyUnit 731 during thewar, based atPingfang inManchuria and commanded by Lieutenant GeneralShirō Ishii. This biological warfare research unit conducted often fatalhuman experiments on prisoners, and produced biological weapons for combat use.[42] Although the Japanese effort lacked the technological sophistication of the American or British programs, it far outstripped them in its widespread application and indiscriminate brutality. Biological weapons were used against Chinese soldiers and civilians in several military campaigns.[43] In 1940, the Japanese Army Air Force bombedNingbo with ceramic bombs full of fleas carrying the bubonic plague.[44] Many of these operations were ineffective due to inefficient delivery systems,[42] although up to 200,000 people may have died.[45] During theZhejiang-Jiangxi Campaign in 1942, around 1,700 Japanese troops died out of a total 10,000 Japanese soldiers who fell ill with disease when their own biological weapons attack rebounded on their own forces.[46][47]
According to historiansBenny Morris andBenjamin Kedar, Israel conducted a biological warfare operation codenamedOperation Cast Thy Bread during the1948 Arab–Israeli War. The Haganah initially used typhoid bacteria to contaminate water wells in newly cleared Arab villages to prevent the population including militiamen from returning. Later, the biological warfare campaign expanded to include Jewish settlements that were in imminent danger of being captured by Arab troops and inhabited Arab towns not slated for capture. There was also plans to expand the biological warfare campaign into other Arab states including Egypt, Lebanon and Syria, but they were not carried out.[51]
Some British soldiers were also poisoned: causing the event to gain international attention.[52]
International restrictions on biological warfare began with the 1925Geneva Protocol, which prohibits the use but not the possession or development of biological and chemical weapons in international armed conflicts.[37][59] Upon ratification of the Geneva Protocol, several countries madereservations regarding its applicability and use in retaliation.[60] Due to these reservations, it was in practice a "no-first-use" agreement only.[61]
The 1972Biological Weapons Convention (BWC) supplements the Geneva Protocol by prohibiting the development, production, acquisition, transfer, stockpiling and use of biological weapons.[6] Having entered into force on 26 March 1975, the BWC was the first multilateral disarmament treaty to ban the production of an entire category of weapons of mass destruction.[6] As of March 2021,183 states have become party to the treaty.[62] The BWC is considered to have established a strong global norm against biological weapons,[63] which is reflected in the treaty's preamble, stating that the use of biological weapons would be "repugnant to the conscience of mankind".[64] The BWC's effectiveness has been limited due to insufficient institutional support and the absence of any formal verification regime to monitor compliance.[65]
In 1985, theAustralia Group was established, a multilateral export control regime of 43 countries aiming to prevent the proliferation of chemical and biological weapons.[66]
Biological weapons are difficult to detect, economical and easy to use, making them appealing to terrorists. The cost of a biological weapon is estimated to be about 0.05 percent the cost of a conventional weapon in order to produce similar numbers of mass casualties per kilometer square.[68] Moreover, their production is very easy as common technology can be used to produce biological warfare agents, like that used in production of vaccines, foods, spray devices, beverages and antibiotics. A major factor in biological warfare that attracts terrorists is that they can easily escape before the government agencies or secret agencies have even started their investigation. This is because the potential organism has an incubation period of 3 to 7 days, after which the results begin to appear, thereby giving terrorists a lead.
A technique called Clustered, Regularly Interspaced, Short Palindromic Repeat (CRISPR-Cas9) is now[when?] so cheap and widely available that scientists fear that amateurs will start experimenting with them. In this technique, a DNA sequence is cut off and replaced with a new sequence, e.g. one that codes for a particular protein, with the intent of modifying an organism's traits. Concerns have emerged regarding do-it-yourself biology research organizations due to their associated risk that a rogue amateur DIY researcher could attempt to develop dangerous bioweapons using genome editing technology.[69]
In 2002, when CNN went through Al-Qaeda's (AQ's) experiments with crude poisons, they found out that AQ had begun planning ricin and cyanide attacks with the help of a loose association of terrorist cells.[70] The associates had infiltrated many countries like Turkey, Italy, Spain, France and others. In 2015, to combat the threat of bioterrorism, a National Blueprint for Biodefense was issued by the Blue-Ribbon Study Panel on Biodefense.[71] Also, 233 potential exposures of select biological agents outside of the primary barriers of the biocontainment in the US were described by the annual report of the Federal Select Agent Program.[72]
Though a verification system can reduce bioterrorism, an employee, or a lone terrorist having adequate knowledge of a bio-technology company's facilities, can cause potential danger by using, without proper oversight and supervision, that company's resources. Moreover, it has been found that about 95% of accidents that have occurred due to low security have been done by employees or those who had a security clearance.[73]
Entomological warfare (EW) is a type of biological warfare that uses insects to attack the enemy. The concept has existed for centuries and research and development have continued into the modern era. EW has been used in battle by Japan and several other nations have developed and been accused of using an entomological warfare program. EW may employ insects in a direct attack or as vectors to deliver abiological agent, such asplague. Essentially, EW exists in three varieties. One type of EW involves infecting insects with apathogen and then dispersing the insects over target areas.[74] The insects then act as avector, infecting any person or animal they might bite. Another type of EW is a direct insect attack against crops; the insect may not be infected with any pathogen but instead represents a threat to agriculture. The final method uses uninfected insects, such as bees or wasps, to directly attack the enemy.[75]
Theoretically, novel approaches in biotechnology, such as synthetic biology could be used in the future to design novel types of biological warfare agents.[76][77][78][79]
Would demonstrate how to render a vaccine ineffective;
Would confer resistance to therapeutically useful antibiotics or antiviral agents;
Would enhance the virulence of a pathogen or render a nonpathogen virulent;
Would increase the transmissibility of a pathogen;
Would alter the host range of a pathogen;
Would enable the evasion of diagnostic/detection tools;
Would enable the weaponization of a biological agent or toxin.
Most of the biosecurity concerns in synthetic biology are focused on the role of DNA synthesis and the risk of producing genetic material of lethal viruses (e.g. 1918 Spanish flu, polio) in the lab.[80][81][82] Recently, theCRISPR/Cas system has emerged as a promising technique for gene editing. It was hailed by The Washington Post as "the most important innovation in the synthetic biology space in nearly 30 years."[83] While other methods take months or years to edit gene sequences, CRISPR speeds that time up to weeks.[6] Due to its ease of use and accessibility, it has raised a number of ethical concerns, especially surrounding its use in the biohacking space.[83][84][85]
Synthetic biology provides the technical capacity to fundamentally alter the bioweapons landscape by enabling the reconstitution of an eradicated or extinct human pathogen. Reports highlight the immediate security concern of "re-creating known pathogen viruses". This capability drastically lowers the barrier to entry for acquiring highly dangerous agents. The deliberate synthesis of the Horsepox virus, an Orthopoxvirus, from commercially acquired DNA segments, stands as a critical academic demonstration of this dual-use capability. This experiment proved that highly complex pox viruses could be engineered.[86][87][88]
Viral Reassortment and Recombination as Dual-Use Risks
Reassortment occurs when two segmented viruses (e.g., influenza, bunyaviruses) co-infect a host cell and exchange entire genome segments. This can generate chimeric viruses with new properties .
Lowen (2018) explains that reassortment "allows exchange of intact genes between related viruses… giving rise to novel genotypes" that may occasionally result in increased viral fitness under selective pressures (Lowen,PLoS Pathogens, 2018).[89]
Recombination involves the joining of nucleic acid sequences from different viral templates into a single genome. This can produce hybrid viruses with traits not present in either parent strain.
Torralba et al. (2024) note that multipartite viruses can reassort even across spatially separated infections, raising concerns aboutunexpected recombinants with enhanced transmission or pathogenicity (Torralba et al.,Virus Evolution, 2024).[90]
Geneious Prime (plasmid design & sequence alignment): Widely used for cloning, primer design, and sequence analysis. Dual-use risk arises from its ability to streamline plasmid construction for pathogenic genes, lowering technical barriers for designing vectors that could express toxins or virulence factors.See: Geneious Prime features overview (Geneious, 2024).[91]
SnapGene (CRISPR guide RNA design): Provides intuitive tools for designing CRISPR/Cas9 edits. While invaluable for therapeutic research, it could be misused to design guide RNAs targeting immune evasion or resistance genes in pathogens.See: Benchling vs SnapGene comparison (OneBrowsing, 2024).[92]
Benchling (cloud-based genetic engineering platform): Enables collaborative design, annotation, and sharing of genetic constructs. Its cloud-based nature raises risks of unauthorized access or covert collaboration for dual-use projects, especially if security controls are weak.See: Benchling platform analysis (OneBrowsing, 2024).[93]
CLC Genomics Workbench (NGS data analysis): Supports large-scale sequencing, variant detection, and metagenomics. Dual-use risk lies in its ability to rapidly identify mutations that enhance virulence or resistance, potentially guiding deliberate engineering.See: Gronvall & Bouri, Biosecurity and Bioterrorism (2008).
PyRosetta (protein structure prediction): Used for modeling protein folding and interactions. Could be misapplied to optimize viral surface proteins for immune escape or host adaptation.See: Chaudhury et al., PLoS ONE (2010).[94]
EpiModel (outbreak simulation): Epidemiological modeling platform for simulating disease spread. While critical for preparedness, it could be exploited to model optimal release strategies for engineered pathogens in a conflict scenario.See: Jenness et al., Journal of Statistical Software (2018).[95]
AlphaFold (protein structure prediction): Breakthrough AI for predicting protein structures. Dual-use risk lies in its potential to predict virulence factor conformations or design proteins that evade host defenses.See: Jumper et al., Nature (2021).[96]
DeepVir (AI for viral transmissibility): Machine learning tool for predicting viral host range, and how contagious a virus can be. Could be misused to optimize viral genomes for cross-species transmission.See: Ren et al., Bioinformatics (2020).
DNA/RNA Synthesizers (e.g., Twist Bioscience): Legitimately used for custom gene fragment synthesis. Dual-use concern: reconstruction of eradicated or high-risk pathogens from sequence data.See: Noyce et al., PLOS ONE (2018) on horsepox synthesis.
Electroporators: Standard lab devices for introducing DNA/RNA into cells. Dual-use risk: facilitating transformation of pathogens with engineered plasmids or synthetic genomes.[97]
Next-Generation Sequencers (e.g., Illumina NovaSeq): Critical for quality control and mutation detection. Dual-use risk: verification of engineered modifications in pathogens, accelerating iterative design cycles.See: Gronvall, Health Security (2017).[98]
Ideal characteristics of a biological agent to be used as a weapon against humans are highinfectivity, highvirulence, non-availability ofvaccines and availability of an effective and efficientdelivery system. Stability of the weaponized agent (the ability of the agent to retain its infectivity and virulence after a prolonged period of storage) may also be desirable, particularly for military applications, and the ease of creating one is often considered. Control of the spread of the agent may be another desired characteristic.
The primary difficulty is not the production of the biological agent, as many biological agents used in weapons can be manufactured relatively quickly, cheaply and easily. Rather, it is the weaponization, storage, and delivery in an effective vehicle to a vulnerable target that pose significant problems.
For example,Bacillus anthracis is considered an effective agent for several reasons. First, it forms hardyspores, perfect for dispersal aerosols. Second, this organism is not considered transmissible from person to person, and thus rarely if ever causes secondary infections. A pulmonary anthrax infection starts with ordinaryinfluenza-like symptoms and progresses to a lethalhemorrhagicmediastinitis within 3–7 days, with a fatality rate that is 90% or higher in untreated patients.[99] Finally, friendly personnel and civilians can be protected with suitableantibiotics.
The United States developed an anti-crop capability during theCold War that used plant diseases (bioherbicides, ormycoherbicides) for destroying enemy agriculture. Biological weapons also target fisheries as well as water-based vegetation. It was believed that the destruction of enemy agriculture on a strategic scale could thwartSino-Soviet aggression in a general war. Diseases such aswheat blast andrice blast were weaponized in aerial spray tanks and cluster bombs for delivery to enemy watersheds in agricultural regions to initiate epiphytotic (epidemics among plants). On the other hand, some sources report that these agents werestockpiled but neverweaponized.[101] When the United States renounced its offensive biological warfare program in 1969 and 1970, the vast majority of its biological arsenal was composed of these plant diseases.[102] Enterotoxins and Mycotoxins were not affected by Nixon's order.
Though herbicides are chemicals, they are often grouped with biological warfare and chemical warfare because they may work in a similar manner asbiotoxins or bioregulators. The Army Biological Laboratory tested each agent and the Army's Technical Escort Unit was responsible for the transport of all chemical, biological, radiological (nuclear) materials.
Biological warfare can also specifically target plants to destroy crops or defoliate vegetation. The United States and Britain discovered plant growth regulators (i.e.,herbicides) during the Second World War, which were then used by the UK in the counterinsurgency operations of theMalayan Emergency. Inspired by the use in Malaysia, the US military effort in theVietnam War included amass dispersal of avariety of herbicides, famouslyAgent Orange, with the aim of destroying farmland and defoliating forests used as cover by theViet Cong.[103] Sri Lanka deployed military defoliants in its prosecution of theEelam War against Tamil insurgents.[104]
During World War I, German saboteurs usedanthrax andglanders to sicken cavalry horses in US and France, sheep in Romania, and livestock in Argentina intended for theEntente forces.[105] One of these German saboteurs wasAnton Dilger. Also, Germany itself became a victim of similar attacks – horses bound for Germany were infected withBurkholderia by French operatives in Switzerland.[106]
During World War II, the US and Canada secretly investigated the use ofrinderpest, a highly lethal disease of cattle, as a bioweapon.[105][107]
In the 1980s Soviet Ministry of Agriculture had successfully developed variants offoot-and-mouth disease, andrinderpest against cows,African swine fever for pigs, andpsittacosis for chickens. These agents were prepared to spray them down from tanks attached to airplanes over hundreds of miles. The secret program was code-named "Ecology".[56]
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In 2010 at The Meeting of the States Parties to the Convention on the Prohibition of the Development, Production and Stockpiling of Bacteriological (Biological) and Toxin Weapons and Their Destruction inGeneva[109]thesanitary epidemiological reconnaissance was suggested as well-tested means for enhancing the monitoring of infections and parasitic agents, for the practical implementation of theInternational Health Regulations (2005). The aim was to prevent and minimize the consequences of natural outbreaks of dangerous infectious diseases as well as the threat of alleged use of biological weapons against BTWC States Parties.
Many countries require their active-dutymilitary personnel to get vaccinated for certain diseases that may potentially be used as a bioweapon such as anthrax, smallpox, and various other vaccines depending on the Area of Operations of the individual military units and commands.[110]
Most classical and modern biological weapons' pathogens can be obtained from a plant or an animal which is naturally infected.[111]
In the largest biological weapons accident known—the anthrax outbreak in Sverdlovsk (nowYekaterinburg) in theSoviet Union in 1979—sheep became ill with anthrax as far as 200 kilometers (120 mi) from the release point of the organism from a military facility in the southeastern portion of the city and still off-limits to visitors today, (seeSverdlovsk Anthrax leak).[112]
Thus, a robust surveillance system involving human clinicians and veterinarians may identify a bioweapons attack early in the course of an epidemic, permitting the prophylaxis of disease in the vast majority of people (and animals) exposed but not yet ill.[113]
For example, in the case of anthrax, it is likely that by 24–36 hours after an attack, some small percentage of individuals (those with the compromised immune system or who had received a large dose of the organism due to proximity to the release point) will become ill with classical symptoms and signs (including a virtually uniquechest X-ray finding, often recognized by public health officials if they receive timely reports).[114] The incubation period for humans is estimated to be about 11.8 days to 12.1 days. This suggested period is the first model that is independently consistent with data from the largest known human outbreak. These projections refine previous estimates of the distribution of early-onset cases after a release and support a recommended 60-day course of prophylactic antibiotic treatment for individuals exposed to low doses of anthrax.[115] By making these data available to local public health officials in real time, most models of anthrax epidemics indicate that more than 80% of an exposed population can receive antibiotic treatment before becoming symptomatic, and thus avoid the moderately high mortality of the disease.[114]
Single cause of a certain disease caused by an uncommon agent, with lack of an epidemiological explanation.
Unusual, rare, genetically engineered strain of an agent.
High morbidity and mortality rates in regards to patients with the same or similar symptoms.
Unusual presentation of the disease.
Unusual geographic or seasonal distribution.
Stable endemic disease, but with an unexplained increase in relevance.
Rare transmission (aerosols, food, water).
No illness presented in people who were/are not exposed to "common ventilation systems (have separate closed ventilation systems) when illness is seen in persons in close proximity who have a common ventilation system."
Different and unexplained diseases coexisting in the same patient without any other explanation.
Rare illness that affects a large, disparate population (respiratory disease might suggest the pathogen or agent was inhaled).
Illness is unusual for a certain population or age-group in which it takes presence.
Unusual trends of death and illness in animal populations, previous to or accompanying illness in humans.
Many affected reaching out for treatment at the same time.
Similar genetic makeup of agents in affected individuals.
Simultaneous collections of similar illness in non-contiguous areas, domestic, or foreign.
An abundance of cases of unexplained diseases and deaths.
The goal ofbiodefense is to integrate the sustained efforts of the national and homeland security, medical, public health, intelligence, diplomatic, and law enforcement communities. Health care providers and public health officers are among the first lines of defense. In some countries private, local, and provincial (state) capabilities are being augmented by and coordinated with federal assets, to provide layered defenses against biological weapon attacks. During thefirst Gulf War the United Nations activated a biological and chemical response team,Task Force Scorpio, to respond to any potential use of weapons of mass destruction on civilians.
The traditional approach toward protecting agriculture, food, and water: focusing on the natural or unintentional introduction of a disease is being strengthened by focused efforts to address current and anticipated future biological weapons threats that may be deliberate, multiple, and repetitive.
The growing threat of biowarfare agents and bioterrorism has led to the development of specific field tools that perform on-the-spot analysis and identification of encountered suspect materials. One such technology, being developed by researchers from theLawrence Livermore National Laboratory (LLNL), employs a "sandwich immunoassay", in which fluorescent dye-labeled antibodies aimed at specificpathogens are attached to silver and gold nanowires.[117]
Researchers atBen Gurion University in Israel are developing a different device called the BioPen, essentially a "Lab-in-a-Pen", which can detect known biological agents in under 20 minutes using an adaptation of theELISA, a similar widely employed immunological technique, that in this case incorporates fiber optics.[119]
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^Barras V, Greub G (June 2014)."History of biological warfare and bioterrorism".Clinical Microbiology and Infection.20 (6):497–502.doi:10.1111/1469-0691.12706.PMID24894605.However, in the light of contemporary knowledge, it remains doubtful whether his hopes were fulfilled, given the fact that the transmission of smallpox through this kind of vector is much less efficient than respiratory transmission, and that Native Americans had been in contact with smallpox >200 years before Ecuyer's trickery, notably during Pizarro's conquest of South America in the 16th century. As a whole, the analysis of the various 'pre-microbiological" attempts at biological warfare illustrate the difficulty of differentiating attempted biological attack from naturally occurring epidemics.
^Medical Aspects of Biological Warfare. Government Printing Office. 2007. p. 3.ISBN978-0-16-087238-9.In retrospect, it is difficult to evaluate the tactical success of Captain Ecuyer's biological attack because smallpox may have been transmitted after other contacts with colonists, as had previously happened in New England and the South. Although scabs from smallpox patients are thought to be of low infectivity as a result of binding of the virus in fibrin metric, and transmission by fomites has been considered inefficient compared with respiratory droplet transmission.
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^Croddy E, Wirtz JJ (2005).Weapons of Mass Destruction. ABC-CLIO. p. 171.ISBN978-1-85109-490-5.
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