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Tiefenbacher",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/10765.jpg",keywords:"Managing Urbanization, Managing Development, Managing Resource Use, Drought Management, Flood Management, Water Quality Monitoring, Air Quality Monitoring, Ecological Monitoring, Modeling Extreme Natural Events, Ecological Restoration, Restoring Environmental Flows, Environmental Management Perspectives",numberOfDownloads:18,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"January 12th 2021",dateEndSecondStepPublish:"February 9th 2021",dateEndThirdStepPublish:"April 10th 2021",dateEndFourthStepPublish:"June 29th 2021",dateEndFifthStepPublish:"August 28th 2021",remainingDaysToSecondStep:"2 months",secondStepPassed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,biosketch:"A geospatial scholar working at the interface of natural and human systems, collaborating internationally on innovative studies about hazards and environmental challenges. 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1. Introduction
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The preamble to the treaty of the Organisation for the Harmonisation of Business Laws in Africa (better known by its French acronym OHADA1 points to the establishment of a new economic order based on the mutual economic benefit of cross border trade. The primary objective of this treaty was to provide a secure legal and judicial environment for business to operate in [1]. This was to be done through the elaboration and adoption of simple modern and common rules adapted to their economies, by setting up appropriate judicial procedures and by encouraging arbitration for the settlement of contractual disputes2. This suggests that uniform laws governing trans-national trade are essential to achieving these goals. To effectively carry out the piece-meal harmonisation of the business laws of member states, specifically through the elaboration of uniform laws, nine Uniform Acts have been adopted till date3. Our interest being sale of goods contract, we shall examine the Uniform Act on General Commercial Law (UAGCL), with respect to its provisions governing sales contracts. Thus, the principal concern of this paper is to critically test the application of the UAGCL to barter-like transactions. The focus on the OHADA business law is to test its functionality in achieving its predictability of business transactions within its contracting states [2].
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The rules on the conformity of goods are not only an integral part of sales law, but also an indispensable obligation by the seller by being inextricably linked to his obligation to deliver the goods. This may explain why ‘goods’ are the very subject matter of a sales contract, and the rules on conformity are what help define this subject matter. Forming part of the economic and legal rights of the buyer, the latter relies on and ascertains the seller’s obligation to deliver the goods in conformity with the contract specifications. This right consists in the delivering of goods of right quality, quantity, description and packaging.
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Consequently, any breach of this obligation would entitle the buyer to establish proof. In this regard, many buyers will base their complaints on defects of conformity of the goods, allege a breach, and invoke remedies. It is important therefore, that there be fairly clear legal rules, particularly those applied by default, that are capable of allocating risks, thereby producing legal certainty and possibly reducing litigation.
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The burden of proof includes the burden of adducing the relevant evidence and the burden of persuasion. The reliance provision is, in other words, an exception to the buyer’s entitlement to the goods fit for a particular purpose, and the burden of proof of the preconditions for that exception lies with the seller. Without these rules, it would often be impossible to say what it is that the seller has agreed to deliver. However, the inevitably broad nature of these rules, together with their considerable conceptual and practical significance, still makes them one of the most frequently litigated issues. All this leads to the conformity rules occupying a central place in any sales contract.
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From the foregoing, the issue of the concept of conformity should be addressed with an open mind because it is dynamic and amorphous. This explains why this study will illustrate that the limits of conformity should not be guided by the distinction between physical and non-physical characteristics of the goods. Other conceptual tools for marking this boundary will be explored in consideration of other parameters irrespective of the intent of the parties as evident in their agreed contract.
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With this in mind, this study seeks in the first place to take a critical look at the rules on conformity in Article 255 of the OHADA Uniform Act on General Commercial Law (UAGCL). The primary objective therefore is to make an expository study of the concept of “conformity” as the basis of the seller’s duty by showing its limits in consideration to other components. It follows with a critical examination of the UAGCL’s approach in allocating the burden of proof of lack of conformity of the goods to the contractual stipulations as agreed by the parties. The study also adopts a critical and analytical approach in interpreting the provisions of the Uniform Act and of foreign instruments regulating sale of goods contracts.
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2. The legal conception of goods
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2.1 Contract of sale of goods
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‘Goods’ form the subject-matter of contracts of sale of goods between the seller and the buyer. The Uniform does not provide any definition to ‘goods’ as the case under the CISG4. Nor is it possible to deduce the meaning of the term by analysing different language versions of the statute. However, from a cursory reading of the Act, a restrictive meaning could be inferred as its provisions basically apply only to moveable tangible goods.
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Reference can be made to the Uniform Act’s Scope and General Provisions. In particular, Article 234 (a) provides that “the provisions of this shall apply to contracts of sale goods”, whereas Articles 235 and 236 restrict the ambit of the Act and by implication, the ambit of ‘goods’. By inference from the restrictions imposed by Article 236 UAGCL, it may be understood that the term ‘goods’ is fairly not extensive, indeed, virtually not all-embracing. It clearly excludes to a greater extent non-physical items, such as electricity, negotiable instruments, and company shares, which are technically ‘things in action’ or incorporeal movables and so are excluded by the plain words of Article 236. Similarly, items of ‘intellectual property’ such as copyrights, patents and trademarks are not corporeal movables and so fall outside the definition, although of course goods may exist, which embody these intellectual property rights
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Even though under the Uniform Act, the notion of ‘goods’ serves to quantify the main obligation of the seller contained in Article 250, which requires that ‘… the seller must deliver the ‘goods’ … as required by the contract and this Uniform Act’, consideration must be given to the type or nature of goods.
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The reason which may be advanced for dismissing intangible or immovable goods from the scope of the OHADA Uniform Act could be that even though these are assets available for trade, they can only be disposed by way of trade or security and not possibly to be transferred physically to another party. The absolute interest in such types of goods may be disposed of outright or may be made the subject of security. Since by virtue of Article 250 para. 1 UAGCL, the main duty of the seller under the contract of sale is to deliver the goods to the buyer, what matters here is the physical transferability of the goods and not necessarily the transfer of a legitimate interest in the goods. The peculiar consideration here does not lie on the identification of the type of goods but rather on the physical segregation and ownership of it. Only in this situation is segregation both possible and necessary to identify the subject of the transfer obligation of the seller under the Uniform Act.
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3. Contracts consisting partially of services
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The UAGCL does not apply to mixed contracts as the case under the CISG, under which the seller provides goods and services. Nevertheless, by inference, these can be treated as unitary contracts rather than separate sales and services contracts [3], and the UAGCL will apply to both parts. This provision further restricts the meaning of ‘goods’, excluding contracts ‘in which the preponderant part of the obligations of the party who furnishes the goods consists in the supply of labour or other services’. The issue of whether software is a good is once again illustrative.
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In modern commerce, an important point, not yet resolved by the OHADA Uniform Act on General Commercial Law, is whether computer software may constitute ‘goods’ within the meaning of Article 234 para. 1. Software is normally embedded in some physical form, such as disks or as part of a package in which it is sold along with computer hardware, that is, computers or computer parts. Therefore, it could be considered as a tangible object capable of it being possible to be transferable. This raises an argument in trying to understand why such an item cannot be considered as a ‘good’ under the Uniform Act. It could without any doubt be covered by the OHADA Uniform Act because such a good is being able to be transferred to another person in a contract of sale in its physical form. Again, there is probability that a disk be physically defective due to a virus for example. In this case, the seller should be liable as the seller of a physically defective car.
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Under French law, goods are known as marchandises. This simply entails a collection of movable assets forming the subject-matter of a contract of sale. This is actually an element of fond de commerce. From this standpoint, there is one clear limit: this meaning will not include any form of immovable property in a contract of sale. Consequently, it can be inferred from the meaning of Article 235 para. 1 UAGCL that it limits the meaning of goods to movable property by its reference to commerçants. The meaning of ‘sale of goods’ limits the very meaning of ‘goods’. Also, this would mean that no sale with a non-trader is of a ‘good’. The general approach that is adopted under the Uniform Act is to apply the OHADA Uniform Act to the commerçant and not to non-commerçant.
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4. The conception and nature of conformity in domestic sales law
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In fact, conformity is a term with a variable content. In English language, conformity is a noun derived from the verb to conform, meaning “agree with” [4, 5]. From this, it becomes clear that the goods should agree with the terms of the contract for them to be in conformity. In other words, the concept of conformity concerns the difference between the object agreed in the contract and that delivered.
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Some particularities of domestic law on the concept of ‘conformity of goods’ in a sale of goods deserves a careful interpretation and understanding [6]. For instance, rules on conformity under the Uniform Act differ considerably from those in common law and civil law. In fact, subtle distinctions can be between the different kinds of defects of conformity under the different laws. Under civil codes5 as well as under the Uniform Act,6 a hidden defect [défaut caché] is distinguished from an apparent defect [vice apparent]; the English Sale of Goods Act (SGA) [1893] distinguishes condition7 from warranties8. Nevertheless, surely merchantability under common law is a similar concept to conformity as the case under UAGCL.
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4.1 Conformity requirements under the Act: Material and functional conformity
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The notion of conformity under the Uniform Act is almost identical to that under the CISG9. The Act views conformity from a dual perspective, that is, material and functional conformity. Whilst material conformity deals with the quality of goods, function conformity on its part focuses on the usefulness of the goods. A recent adoption by the Uniform Act is the condition that goods must conform both materially and functionally before they are judged acceptable by the Act. As such, sellers have to respect this prescription before delivering their goods10 [7, 8].
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The conditions of quantity, quality, description, packaging, particular purpose, and sample are encapsulated into the concept of conformity in the Uniform Act as contained in its Article 255 [9]. Article 255 thus states as follows:
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“The seller shall deliver the goods according to the quantity, quality, specification, and packaging provided for in the contract. Where the contract is silent, the seller shall deliver goods in conformity with the purposes for which goods of that nature are generally used, and the goods must match the sample or model which was presented to the buyer by the seller. The seller also must deliver the goods that are packaged according to the usual method of packaging goods of the same nature or failing which, in a manner to ensure their conservation, and protection”11.
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These implied that conditions deserve careful treatment because of the protection that they now offer the buyer of goods, who is almost invariably in a weaker position than the seller. This can be explained by the fact that, most of the times, the seller seems to be the manufacturers of the goods. As a result, these terms protect buyers’ interest since they ensure that the purchased goods are neither deficient nor defective. The Act thus makes it clear that any breach of the provisions by either parties shall be interpreted as non-respect of terms of contract. It is important to state that the terms of a contract, which include conditions and warranties, of sale of goods could be implied or expressly stated. It is now clear that the common law principle of caveat emptor which used to focus on the buyer now places emphasis on the seller’s awareness (caveat venditor) [10]. Implied conditions and warranties are not stated by the parties during negotiations or included in a contractual document, but nevertheless form part of the contractual provisions.
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Implied terms as implied by law are geared to ensuring a minimum of business efficacy12, regardless of the parties’ paramount intention to create a workable contractual agreement13. Some contracts of sale are very detailed; the parties deal with all or most eventualities. However, in others, the only element that the parties deal with is identifying the goods to be sold and the price to be paid.
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The seller is obliged to be conversant with aspects of quality, quantity, specification, and the packaging of goods, which falls within material conformity as well as features linked to functional conformity such as the suitability, purpose, or purposes and specificities of the usefulness of goods of similar design.
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Material conformity therefore consists of four elements derived from the contract: quantity, quality, description, and packaging.
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4.2 An inquiry into the nature of lack of conformity
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4.2.1 Apparent defects
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Article 258 of the Uniform Act raises no difficulty as to defects which would have been apparent on a reasonable examination of the goods by the buyer immediately after delivery. In practical terms, examination by the buyer of any apparent defects to ascertain the seller’s obligation operates from the moment delivery has been effected14. After this exercise, the buyer must give notice of a lack of conformity discovered within 1 month from the date of delivery. If he fails to observe these requirements, the buyer will be deprived of his right to claim redress for non-conformity.
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4.2.2 Hidden defects
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Moreover, it is advisable that the buyer declares any detected defect observed in a purchased item within a year to enable the seller to make up for any deficiency within his/her competence15. The period of 1 year is enough time for the buyer to notice hidden defect in a purchased good. The foregoing condition notwithstanding, it is necessary to state that the detecting of defect goes beyond according to a timeframe since some goods require immediate inspection and adjustment even before they are sold and bought. In this case, therefore, the buyer gets to understand any hidden defect before the commercial transaction.
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However, the detecting of hidden defects is difficult, especially with frozen and canned/tinned goods that are always meticulously packaged. In such cases, there is little or no opportunity for instant or early detection of defects.
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The problem raises two complex issues. The first is how to establish the liability of a seller or retailer who sells food products in tin or other sealed container because of the difficulty to identify any injury caused to his customers either from some foreign substance in the can or from unwholesomeness of the food and the second is how to establish the liability of a seller who sells goods with some defects in bulk. In this situation, the seller may not be faulted for such hidden defects owing to the difficulty involved in detecting such defects in bulk goods. These complex situations account for the emphasis of Uniform Act, in relation quality goods, that the seller shall be liable to deliver the goods “in the usual manner” that goods are packaged16.
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Examples of such nature regarding the determination of hidden defects in goods can never solve all the problems related to quality. Nevertheless, with the aid of Article 255(2) of the Uniform Act, the intent of the parties can be construed in their agreement. In striving to achieve this purpose, two questions could guide this exercise: The understanding of the contractual provisions defining quality goods as spelt out by Article 255(2) and the usefulness of the goods are two basic conditions that could guide the parties, as the strive to satisfy their intent. An understanding of these conditions would certainly curb misunderstanding with regard to fitness of ‘ordinary’ use of goods as stated in the contract.
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It follows therefore that the nature of the kind of non-conformity that the buyer is expected to reveal are two, namely apparent and hidden defects. This type of test is likely to pose challenges involving goods of varying grades.
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4.3 Different quality test: a matter of interpretation
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A further inquiry into the notion of quality is necessary. This discussion points to the various quality tests the buyer may alleged non-conformity of goods. In fact, this raises a debate in trying to clear the confusion surrounding the basis of the seller’s liability for the non-conformity of goods. This discussion, in turn, makes it necessary to choose between various quality tests.
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Against this background, the Act’s default rule in Article 255 para 2 appears narrow and limited in its content and scope. On its face, this provision does not rely on any notion of quality, with the only relevant question seemingly being whether the goods are fit for “the purposes for which goods of the same description would ordinarily be used”. In other words, the Uniform Act only seems concerned with whether the goods are fit for their ordinary purposes and not with quality. Rather, quality is a broader notion that may include not only fitness for ordinary purposes, but a number of other aspects such as the goods’ physical state and condition, intrinsic qualities and features, safety, durability, appearance, finish, and freedom from minor defects17. It is an undeniable fact that these other aspects of quality could be attributable to the dynamic decisions of individuals and group choices and satisfaction based on information gathering relating to a particular commodity [11]. Further exploration of the problem centres on the considerable evidence that quality variation is greater in group unanimity than in personal preferences over certain goods. Here, the commercial buyer may be inclined to conform and to choose goods required by group customers in the market.
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As a general rule under the Uniform Act, conformity of the quality of the goods will be met if and only if the usage criterion is also satisfied18. In fact, the commercial utility of the goods seems to be the guiding rule to the commercial buyer under the Uniform Law in ascertaining the seller’s responsibility. This can be best explained by the fact that the utmost preference to the commercial utility of the goods by the commercial is guided by several different ways in which interpersonal influences impact variety-seeking behaviour of consumers [12]. In fact, changing social habits may require merchants to select a variety of items appropriate to the demands of divergent contexts and audiences with the view of acting in conformity with consumers’ preferences. Understandably, consumers’ individual choices tend to conform to or diverge from the choice of another group of consumers [13]. It follows therefore that the material conformity of the goods to the commercial buyer is guided by individual and divergent preferences of its end users. The result of the non-respect of both material and functional quality or either of the qualities, often insinuates a breach of contract. In a case where the quality is deemed essential, the breach may result in a fundamental breach with the consequence that the buyer may either ignore the contract or request the total substitution of the purchased good\n19.
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Conversely, many domestic legal systems have used varied notions in ascertaining the ultimate default rule of conformity of the goods in order to safeguard the buyer’s satisfaction. Notions of quality such as “average, “merchantable,” “acceptable, or “satisfactory quality” have been the measuring yardsticks meanwhile “fitness for an ordinary or a common purpose” is merely one of the components referred to in some legal systems to make a sound judgement on the notion of quality or to know whether the goods meet the required standard of quality20.
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The above approach can be said to be associated with the merchantability test, which obtains with some common law systems. This test is viewed as comprising more than just the saleability of the goods as those terms in the parties’ contract (with description, fitness for purpose, and acceptance being other relevant aspects of merchantability) [14].
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4.3.1 Ethical values
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Ethical value is an emerging applied subject. It is still currently debated as it is applied to all aspects of business conduct, the manufacturing of goods, and the quality standard of goods. In general, ethical values appear to contribute to the development of legal systems in which contracts are enforced fairly, and bribery and corruption are less prevalent, enabling business entities to have equal access to legal process and equal protection under the law.
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At the international level for instance, private initiatives such as the principles laid down in the United Nations Global Compact covering issues such as human rights, labour, environment and anti-corruption aim to assist its member countries to comply with the international standards required on these issues. In Africa, ethical values are known and recognised in international and national sales contracts as one of the solutions to the defective goods as from the manufacturing process.
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An ethical value contractual clause can be express or implied in the contract of sale. When implied, they may be inferred from trade and usages. Whether expressly or impliedly in a contract of sale, they become part of the contract and may be enforced, or their violation sanctioned, in the same way as with any other term. However, the parties agree on their express stipulation [15].
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In Cameroon, for instance, most organised professions and some trade associations have codes of practice which set out ethical behaviour for members in their dealings with their clientele21. For instance, the company in charge of distributing energy makes available to the public an e-mail address for anyone who wants to make a complaint and report any violations of their Code of Business Conduct and Ethics. A clause ensuring ethical code and conduct is prior inserted in the contract signed by customers before they benefit from the services of the company. This is done in favour of its customers.
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In line with the above example, it should be pointed out that in most, almost all contracts of sales of goods, including both domestic or international contracts, the quality of goods is not restricted to its physical appearance; it also has to comply with any other requirements which are prescribed by the law or subject to trade and usages22. Therefore, if the origin of the goods prescribes an ethical value standard at production, the standard will definitely constitute a quality of the goods. As a result, the requirement of quality will not be met if the goods produced do not meet with the ethical standard as required. It could therefore be agreed that people incorporate more variety into their consumption decisions when their behaviour is subject to public scrutiny [16].
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It is not easy to determine the non-conformity based on the ethical value23. Such difficulty is due to the hidden character of its nature and also to the fact that it does not physically influence the physical feature of the goods. If the breaching party does not of good faith disclose or cure the defect, an expert may sometimes be required to determine the breach.
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Unless the parties have agreed otherwise, failure from the seller to deliver the goods in conformity with the ethical standards amounts to a breach of contract. In particular, when the seller is required to deliver goods that are fit for the purpose for which goods of that nature are commonly used or when a special request on an ethic (such as health and environmental considerations) was made known to the seller24. The buyer remedy will be its rights of redress25 and claim for damages26.
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4.3.2 Good manufacturing practice
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GMP refers to the “Good Manufacturing Practice”. This is a well-known regulation whose purpose is to ensure the appropriate manufacturing process that would impact on the quality of the products. Good Manufacturing Practice is a series of regulations that are binding on the contracting business parties, (is agreed upon) requiring food producers, manufacturers, pharmaceutical companies, and packagers of drugs to ensure that their products are safe, pure, and totally effective. These regulations are then meant to ensure a level of quality approach to manufacturers of goods in order to protect consumers from contracting for non-effective and dangerous goods. In summary, Good Manufacturing Practice plays a paramount role of quality assurance that ensures that products are consistently produced and controlled to the quality standards appropriate for their intended use and their legal requirements. In some cases, it is simply required by the marketing board or in some countries by the government department in charge of regulating trade and commerce27.
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Sometimes, GMP is referred to as Cgmp- meaning “Current Good Manufacturing Practice”. This is a mere call to producers, packagers, or manufacturers to update their machineries, systems or equipment in order to comply with the new technology, capacity and /or legal requirement in their business. Generally, GMP’s guidelines provide regulation on products such as foods, drugs, blood and medical devices. Accordingly, countries do update their cGMP regulations for local companies to act in accordance with the legislation. In Cameroon, for instance, the Medical Council is a statutory body that was established in terms of the medicines and related substances control to oversee the regulation of medicines. Nonetheless, the “Standard and Quality Agency (ANOR)” and the “Service of Norms” in the Ministry of Trade and Commerce have a wide mandate to ensure the quality of products in meeting the required standards28.
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Even though not prescribed by the UAGCL, such a business practice or measure is relevant in enhancing the warranty of quality of goods that flood the markets. This goes a long way to improve on the quality of goods which the end users would obtain from commercial buyers. Respect of such a measure is left at the wishes of the parties in a contract of sale or could be implied from the trade and usages of specific goods in a given branch of business. This is the very meaning of Articles 237 and 239 of the UAGCL.
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5. Conformity and other related concepts as characteristics of the ‘goods’
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Generally, goods have certain defined physical characteristics which in everyday business practice the parties have full knowledge of when contracting. In some cases, certain states of affairs may be so inextricably linked to the goods as to be ‘non-physical characteristics’ of the goods. In this regard, the parties may in concluding their sale insist on some features connected to the goods but are not physically part of the goods. These could concern the tangible and intangible things connected to the good. Nevertheless, the dichotomy between intrinsic and extrinsic characteristics of a ‘good’ is fundamental in determining the functionality of the goods. In effect, these characteristics are broad and can be described in various terms, for example, intrinsic/extrinsic; embodied/unembodied; tangible/intangible; corporeal/incorporeal; objective/subjective; real/intellectual and physical/non-physical.
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A key example of non-physical characteristics that the parties may view as part of the goods emerges from the field of marketing. In that field, ‘[the idea of tangible and intangible attributes [of goods] is well established’ [17]. Tangible characteristics are those that ‘are physically present or can be seen, experienced or measured in some way’. Intangible characteristics, such as reputation, quality image, and country of origin, are ‘understood using cognitive processes and also often contain an emotional dimension’29. It should be stressed that these intangible characteristics most impact the consumer behaviour in guiding their choices of goods they buy. It, therefore, ascertains the economic value of goods30. Hence, these non-physical attributes are conceived of as part of the goods.
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5.1 Brand
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A brand of a good is an exemplified example to be conceived as a non-physical attribute of the goods. By definition, a ‘brand’ is a commercial concept that describes’ the impression of a product in the minds of potential users or consumers’. This explains the more reason why the wide sale of certain goods in some international markets is due to the strength of their brand and reputation publicised by mass advertisements and persistently demanded by a group of customers. That is why it is common today to find consumers in places across the world wearing, driving and drinking the same brands of certain goods [18]. This is considerable evidence that quality variation is greater in group unanimity than in personal preferences over certain goods [19]. This is another clear indication of the fact the commercial buyer of goods in her business will be seeking to conform to group norms.
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Brand is conceived as an attribute of the goods, rather than something external to them31. Related to this is the notion of ‘brand equity’. The brand equity in a good is measured by the difference between the inherent value of the good and the perceived value (or market value) of the good. Another justification for this is the public habits and beliefs in certain virtues of these goods, which have stood their test of time. Such a picture is typical in Cameroon where from socio-economic and psychological perspectives, the purchasing public still hold tight to the buying of some goods of old reputable trademark or taste. For example, certain brand of goods such as, shoes like “pierre cadin” from France, “clarks” from England are still believed to be durable even sold as used goods. The sale of used goods or second-hand goods is an everyday practice where is visible to find a line-up of stores in Cameroonian cities such as Douala and Yaoundé operating in this trade. Goods vary from cars, household equipment to consumer goods. Based on observations from field trips conducted by the researcher in the above mentioned cities, it was gathered that consumers prefer certain used goods irrespective of their diminished quality and safety but rather simply because of their cheap prices, accessibility and long standing brand [20].
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The law recognises that reputation and customer awareness are worthy of protection by allowing the enforcement of intellectual property rights. Although it is not a legal concept, brand equity has been referred to as being ‘[a]t the core of a trademark’s value and a manifestation of the legal concept of trademark good will’ [21].
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The branding and reputation of the goods are connected to intellectual property in the goods. The seller’s duties in respect of conformance and intellectual property rights are interrelated rather than mutually exclusive. Certain characteristics of the goods may indeed be relevant to both duties. As will be demonstrated, the UAGCL’s provisions relating to intellectual property are apt to deal only with the existence of certain non-physical characteristics of the goods. The content of those characteristics must be dealt with under the principle of conformity. The seller has express obligations in relation to trademarks and other intellectual.
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property rights under Article 260(1) UAGCL: the seller shall deliver the goods with the assurance that no third party has a right or claim to them, unless the buyer accepts to collect the goods under such conditions.\n
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From a reading of Article 275 UAGCL, it is clear that the main purpose and effect of the seller’s duty to deliver is to require the seller to transfer the property or title to the goods to the buyer. This implies that the seller should not necessarily be the owner of the goods. The fundamental requirement here is to qualify the seller’s right to sell and also the third party’s right(s) over the goods. As concerns the seller’s right to sell, the seller must not necessarily be in the possession of the goods and be able to transfer property to the buyer. To avoid any third party’s claim, the seller must be the right holder of some patent, trademark or other proprietary interest over the goods. Any possibility for the third party to have access to the goods on such reasons will amount to a defect in title and the seller will be liable for this. The essence here is for the seller to be able to transfer to the buyer a title overriding that of the true owner.
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6. Burden of proof for the non-conformity of the goods with the standard
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As a general principle, the allocation of the burden of proof under the Act, seeks to know whether or not the goods are presently in conformity with the applicable standard as contained in Article 255. This mostly guided by the principle of proof proximity, which states that conformity or non-conformity of any good is determined by the individual who is in possession of the said good. This means that when a good is still with the seller, he/she is in the rightful position to establish and prove that the said good conforms or not. By the same token, once the buyer purchases the good, he/she assumes the responsibility of establishing conformity or non-conformity of the good in the case of an incident.\n
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One relevant general principle under the UAGCL is that the assessment of the conformity of goods sold starts upon delivery32. In fact, it is understandable that it is the buyer’s obligation to examine the goods in view to reveal any lack of conformity. Therefore, the law grants an opportunity to the buyer to inspect the goods and report to the seller whether or not the goods are in conformity with the contractual obligation as agreed upon with the seller [22]. The question that arises is whether the buyer’s knowledge, usually derived from having an opportunity of pre-contractual examination of the goods, should be relevant to deciding what conformity obligations were imposed on the seller by the contract and, even if not, whether the facts of the buyer’s inspection or assessment is hardly relevant to determine the content of the seller’s obligation33.
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The philosophy behind the application of Article 255 UAGCL directs that a buyer ought to be able to rely on the agreed arrangements. Therefore, if there is any discrepancy between the contract and the goods inspected, the buyer has a right to cause the seller to redress the situation. This applies regardless of whether the buyer knows of a defect, or if it was agreed that the seller shall deliver the goods without defects [23]. At this point, the parties must be assumed to have expressed their wishes in their written agreement in relation to the requirements for the goods in question. This means that an analogous or expanded interpretation of the provision so as to apply it to Article 255 para. 1 of the Uniform Act seems in principle to be a restricted application of the caveat emptor principle.
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When reference is made by the contracting parties, what is normally meant is inspection by the buyer. However, control of quality may also be effected by the seller, in which case its quality certificate is attached to other documents (which is significant in respect of certain goods since quality may be considerably affected during transport by reason of environmental and other circumstances). This method of inspection is provided by some governmental or international bodies. The certificate of inspection is the only proof of what it is bound to furnish under the contract. There is, however, another view to the effect that the other party is always entitled to a further inspection34.
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The answers to these preoccupations exert an influence on contract interpretation depending on the particular circumstances. Presumably, these points are too obvious for the drafters of the Uniform Act not to have been aware of them considering the fact that Article 255 para. 1 does not concern the terms implied by default but concerns what the contract itself provides. This clearly shows the limits of the application of the concept of conformity under the Act which presumably seems to be detrimental to the business climate. In actual fact, in addition to that standards contained in the contractual stipulations, other national and international norms of goods need to be considered. Rapid technological changes with a corresponding impact on consumer behaviour is taking place within an increasingly liberalised marketplace in which the global drive to compete brings new challenges to regulatory authorities in setting up conformity standards of goods.
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From the above strand of reasoning, it is certain that irrespective of the contractual stipulations, it is also of prime importance to note that the commercial buyer could dictate other requirements to the seller as driven by the desires of consumers. In addition, there is also necessity of the seller to have knowledge of surrounding facts requiring the goods to conform with certain local and international standards. Nevertheless, no matter the argument, the buyer’s evidence as to any lack of conformity can only be validated through his duty to examine as well as supported by any information which was brought to him by the seller.
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6.1 Burden of proof for the non-conformity of the goods at the time risk passes
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In real commercial practice, there is usually the practical difficulty in ascertaining the allocation of the burden of proof with the purpose of establishing a defect of conformity of the goods which already existed at the time risk passed, or existed before that time. The situation under the Act is ambiguous in that there is no clarity with regard to the charge on the seller who deals in goods that do not conform or are observed to have hidden defects. Moreover, the Act does not clearly spell out the seller’s responsibility when his/her goods are defective or do not conform as a result of a particular kind. Arguably, the notion of conformity under the Act falls within the meaning of the subjective understanding of a “defect” [24].
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As an evidence to the forgoing fact, the purport of Article 256 UAGCL which suggests the fact that a buyer can only make a case for any lack of conformity after the risk passes irrespective of the nature of the defect. In fact, the strong implication is that the burden of proof has shifted onto the buyer in order to establish the seller’s liability when delivery has duly been effected. Most often when the buyer verifies a good to be purchased, the intension is hardly to establish its actual state; rather he/she tries to determine the good’s non-conformity aspects in order to demand immediate reparation. The new Uniform Act has further empowered the buyer to notify the seller of any observable defect. Despite this responsibility given to the buyer, the nature or kind of defect the buyer is supposed to report to the seller is not clearly defined.
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There remains, however, a serious problem about the kind of defects which the buyer is expected to report to the seller. Nevertheless, a clear right of action has been imposed unto the buyer to establish the seller’s responsibility for apparent defects and for hidden defects35 discovered by the commercial buyer particularly when delivery has taken place.
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At this juncture, it would be necessary to make an inquiry into the type of risk that can be transferred or passed to the buyer after delivery before examining the kind of defects the buyer is expected to reveal.
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6.1.1 Which risks are transferred to the buyer?
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Risk of “loss” in a sales context refers to the allocation of financial responsibility for the injury or destruction of goods that occurs whilst the goods are changing hands from seller to buyer [25]. To identify the appropriate moment for passage of this risk is the task of the law governing risk of loss. That law is currently found in Article 277 et seq. of the Uniform Act. The Uniform Act contains no definition of the types of risks governed by the rules on transfer of risk, thus leaving uncertainty. First, one must look at the risks that fall within the scope of the Act36.
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6.1.1.1 Liability for loss or damage to the goods
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The wording used in the Uniform Act is “loss or deterioration”37 contained in the Article 277 para. 2. Physical risks to the goods including their destruction are covered by the concept of “loss”. The Uniform Act’s risk-of-loss rules clearly limit their ambit to loss or damage to “the goods” (that is, the goods sold) or “in respect of goods sold”. By analogy, the causes of loss could be broad encompassing disappearance of the goods, including theft, misplacing the goods, their transfer to a wrong address or person, and mixing up the goods with other goods are included.
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This situation was vivid during a field trip conducted by the researcher in markets of some big cities in Cameroon: Douala and Yaoundé38. Most big merchant-sellers of second- hand goods like shoes for example find in their bulk of goods damaged shoes and because the risk has already been transferred to them by their supplier in their country, such as Dubai and China, they are unable to return the goods. Therefore, because of financial interest, they are forced to sell them at relatively cheap prices.
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Another concern is the documents pertaining to the goods. The risk-of-loss rules of the Uniform Act apply as easily to documents as to goods. This study rather holds strong to the fact that the risk -of- loss of a document connected to the goods, should simultaneously pass together with the risk for the goods. The time and place to hand documents may at times not reach the buyer concurrently. In a case where there exists no agreement to this effect, the delivery of documents may be expected just in time for their use, for taking delivery of the goods or for their import in accordance with the trade usage39. Thus, if the documents are lost before they are delivered; the risk would not be treated similarly as for goods. Consequently, the holding of relevant documents of the goods by the seller does not affect the passing of risk40. The buyer could rather be contended to claim remedies for non-conforming delivery of documents, by applying to the courts for avoidance of the contract, which would stop risk relating to those documents from passing41.
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7. Proof as matter regulated under the uniform Act
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Considering that Article 255 of the Act neither states the liability attached to proof nor points out on whom the responsibility of determining the conformity and non-conformity standard of a good as well as the timeframe of declaring such information, UAGCL principle have to be applied to establish the seller’s liability in a case where a purchased good does not conform with acceptable standards.
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Nonetheless, one relevant general principle under the UAGCL is that a party who asserts a right must prove the necessary preconditions for the existence of that right. This is clearly evident under Articles 258 and 259 para. 1, which gives the burden to the buyer to notify the seller cases of lack of conformity. The buyer’s right to do so is to ensure the seller’s obligation in delivering goods in conformity with the express contract terms pursuant to Article 255 para. 1 of the UAGCL. It suggests that the burden of proof includes the burden of adducing the relevant evidence and the burden of persuasion. This comes into play by the reliance of the buyer on the seller’s conformity obligation. In other words, an exception to the buyer’s expectation to the goods fit for a particular purpose, and the burden of proof of the preconditions for that exception lies with the seller. However, if the seller does not raise the issue of reliance, the goods’ fitness under Article 255 para. 2 will be presumed.
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The “rule and exception” principle of the allocation of burden of proof may not always be applied strictly in practice because the burden of adducing evidence is sometimes placed on a party who simply has better access to evidence but who would not otherwise bear this burden on strict principles of the allocation of burden of proof42.
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From the above, as well as in line with Articles258 and 259 para. 1, it can be deduced that the buyer has to prove the factual prerequisites of the provisions upon which he wants to rely for its claim or defence. It follows from the above that in absence of an explicit regulation in the UAGCL, the allocation of the burden of proof in relation to the various factual requirements in establishing the seller’s liability for non-conforming goods has to be exercised primarily on the basis of the general principles underlying the UAGCL.
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These general principles are to be found first of all in the few provisions which explicitly address the question of burden of proof, in particular Article 294(1). It states:
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\nA party is excused from his duty to render performance if he can prove that it is made impracticable without his fault by the occurrence of an event, namely; due to a third party or the occurrence of a force majeure. A force majeure entails events which happen beyond the party’s control and which could not be reasonably foreseeable. However, there is no exemption if the failure to perform has been caused by a third party appointed by the defaulting party to perform all or part of his contractual obligations\n43.
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Inherent in these general statements of excuse are four elements: (1) performance has become impracticable; (2) the non-occurrence of the cause of impracticability was due to force majeure; (3) the party asserting the excuse is without fault; and (4) the party seeking excuse did not assume greater obligations in the contract.
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On the strength of this, as well as in accordance with Articles 235 (a) and 282 UAGCL it can be deduced that each party is under the obligation to prove the factual prerequisites of the provisions as the basis of his/her complaint and justification. This situation originates from Roman practice expressed in Latin as ei incumbit probatio, qui dicit non qui negator actori incumbit probatio; translated in English as ‘rule and exception-principle.’ It is important to underline that the respect of equity is the driving force behind this rule, since both the UAGCL and the CISG do not expressly elaborate on the principle of proof proximity44.
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This rule is supplemented or modified by considerations of equity according to which each party has to prove those facts which originate from its sphere. The basis for this principle proof proximity in the UAGCL as in the CISG is less clear [26, 27]. Consequently, the courts will presumably generally limit themselves in stating the existence of the principle without giving any further justification. The proof proximity pays attention to the ability of the party to gather evidence as well as the relevant facts and issues presented as proof. This implies that once a good is delivered to a buyer without a prior indication of deficiency from him/her (the purchaser), the responsibility to prove any claim based on lack of conformity of the delivered good becomes that of the buyer. This seems to be the spirit surrounding the provision of Article 256 of UAGCL. It states:
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\nConformity of the goods shall be appraised as of the day of delivery, even if defects appear only later\n45.
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The rules on conformity under the UAGCL are by no means an exception in dealing with issues of proof as the case under the CISG. The point to be addressed here in the first place is the burden of proof. The burden of proof is not a legal obligation, but by its legal nature, a duty. The duty represents an obligation to oneself and not to the other party in a contract. The duty to proof is closely connected with the buyer’s duty to examine the goods and to notify the seller. Namely, the seller will be liable for the non-conformity of delivered goods only if the buyer gives notice pursuant to Article of the UAGCL.
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The main purpose of the examination is to determine whether or not the goods are in conformity with the contract, that is, to reveal defects in quality, quantity, description and packaging. In fact, it is only on the result of the examination that the buyer can make a claim for nonconformity. It follows therefore that, burden of proving non-conformity rests on the buyer.
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8. Overview of the various allocation of burden of proof in practice
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It is expressly clear from the provisions of the Uniform Act that the allocation of the burden of proof for the seller’s liability for non-conformity of the goods at the time the risk passes is imposed on the buyer. It is however rare to find under the Act an apportion of the burden to proof to the seller. There is a lack of necessary specificity and distinction to attribute it clearly to the seller.
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In light of that and other considerations, attention should also be given to the fact that the Act is completely silent on the procedure for establishing the burden of proof. This can be explained by the fact that such an issue which is beyond the OHADA’s scope of application and consequently be governed by the non-harmonised national laws of member countries. Consequently, such a question must be left to the courts of member states as a matter of procedural law. In addition, it is inappropriate for the Act, which relates to the cross-border sale of goods, to deal with matters of evidence or procedure. In fact, they are still some hurdles that beset the uniform working of the OHADA Uniform Act on General Commercial, considering the glaring differences between the procedural and evidential laws operating in the member States on matters of proof. This is as a result of the prevailing differences in legal cultures, procedures in the area of criminal proceedings which is out of the scope of the Act. Therefore, a degree of non-uniformity can be expected in matters of taking evidence and more broadly in allocating burden of proof.
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In fact, the allocation of the burden of proof under the Uniform Act rests primarily on the basis of the actori incumbit probatio principle. Thus, the burden of proof is largely dependent on the position of the parties in the process, that is, who invokes Article 255 in its favour.
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The common practice is that the responsibility of proof is often on the basis of proof proximity principle, whereby the responsibility to prove the non-conformity with standards of a good is transferred from the seller to the buyer once a purchased item is delivered to the buyer. The implication here is that the seller is responsible for proving the conformity or non-conformity of his/her good only a purchaser has not yet taken possession of or demanded the reservation of the particular good. Therefore, the moment a buyer receives the delivered good raising the question of non-conformity, any further proof non-conformity would be his/her responsibility.
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It is suggested, however, that legal predictability should not be undermined any further by the introduction of the proof proximity principle into the UAGCL. As already alluded to, proof proximity can easily contravene the rule and exception principle, and its introduction necessitates a choice between the two, either as a matter of general principle or in the particular case. That, in turn, gives rise to an additional layer of complexity and unpredictability.
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Reaching a substantial degree of international agreement on the rule and exception principle is a hard-earned achievement, which has potential to promote legal certainty in all areas falling within the Act’s scope. From this standpoint, recognising proof proximity as the Act’s general principle would be an unwelcome development.
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9. Conclusion
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The seller’s duty in exercising his material duty of conformity under the UAGCL is fraught with some difficulties. One of such difficulty is to establish the seller’s liability for the non-conformity of goods as to the specifications of a contract of sale of goods. There is still some confusion and uncertainty regarding the notion of conformity under the Act because the concept of conformity is dynamic and ambivalent. There is really a need to search for a quality standard, which can underpin the ultimate default of the fitness for ordinary purposes and engulf other aspects of conformity. This will greatly be in the interest of the commercial buyer in order to conform with the demands of his consumer buyers.
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The seller is equally expected to deliver goods which will satisfactorily serve the purpose for which commercial buyers intend to use them. This suggests therefore that the seller’s expected obligation to deliver the adequate requirements to the goods under the contract would hardly in practice be that. This therefore raises the confusion and uncertainty in determining the seller’s liability for the non-conformity of goods as to the specifications of a contract of sale of goods under the UAGCL. The concept of conformity must be handled in consideration with a number of issues, irrespective of the contractual stipulations agreed by the contracting parties.
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It has also been evident that the issues of proof of lack of conformity under the Act be useful to introduce an overarching quality standard. However, to what extent should a sales law instrument govern the matters of proof, such as burden and standard of proof, and the evaluation and admissibility of evidence? It has been argued that the UAGCL is not capable of dealing adequately with the admissibility of evidence. Consequently, this may have considerable impact in establishing the seller’s liability and on substantive law rights. No consistency in the exercise of this right is available under the UAGCL. There are undeniably differences in legal cultures, procedural environments, and views of the purpose of judicial proceedings in the member states.
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To conclude, there is little doubt that the UAGCL has proved to be capable of resolving the issue of conformity of the goods. Attention should be given in drafting new provisions on conformity. Finally, to what extent should the UAGCL govern the matters of proof, such as burden and standard of proof, and the evaluation and admissibility of evidence. This calls for a fundamental harmonisation of the adjectival laws on matters of proof in order to maintain a higher degree of the uniform character of the OHADA laws.
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\n\n',keywords:"sales, goods, legal, discourse",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/73027.pdf",chapterXML:"https://mts.intechopen.com/source/xml/73027.xml",downloadPdfUrl:"/chapter/pdf-download/73027",previewPdfUrl:"/chapter/pdf-preview/73027",totalDownloads:185,totalViews:0,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,dateSubmitted:"November 26th 2019",dateReviewed:"May 26th 2020",datePrePublished:"August 28th 2020",datePublished:"December 23rd 2020",dateFinished:"August 20th 2020",readingETA:"0",abstract:"This chapter is primarily concerned with the fact that the concept of conformity is dynamic and amorphous as it is recognised as an impetus to economic development and plays a major role in matters of sale of goods within an economy. In making an assessment of the seller’s duty of conformity to a contract of sale of goods as governed by the OHADA Uniform Act on General Commercial Law, this study argues that the concept of conformity is limited rather than broad that should appropriately encapsulate the physical and non-physical things that could form the object of a contract of sale. It therefore explores other aspects that could be considered as part of the ‘goods’ for the purposes of the conformance duty in establishing the limits of the seller’s liability. Thus, adopting an empirical and in-depth analysis of primary and secondary data, this study therefore holds that the question of conformity of goods can conveniently be addressed from a number of different angles: contract law, consumer patterns, local and international standards, and the principles of caveat venditor and caveat emptor.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/73027",risUrl:"/chapter/ris/73027",book:{slug:"banking-and-finance"},signatures:"Djieufack Roland",authors:[{id:"316005",title:"Dr.",name:"Djieufack",middleName:null,surname:"Roland",fullName:"Djieufack Roland",slug:"djieufack-roland",email:"orolly@yahoo.fr",position:null,institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. The legal conception of goods",level:"1"},{id:"sec_2_2",title:"2.1 Contract of sale of goods",level:"2"},{id:"sec_4",title:"3. Contracts consisting partially of services",level:"1"},{id:"sec_5",title:"4. The conception and nature of conformity in domestic sales law",level:"1"},{id:"sec_5_2",title:"4.1 Conformity requirements under the Act: Material and functional conformity",level:"2"},{id:"sec_6_2",title:"4.2 An inquiry into the nature of lack of conformity",level:"2"},{id:"sec_6_3",title:"4.2.1 Apparent defects",level:"3"},{id:"sec_7_3",title:"4.2.2 Hidden defects",level:"3"},{id:"sec_9_2",title:"4.3 Different quality test: a matter of interpretation",level:"2"},{id:"sec_9_3",title:"4.3.1 Ethical values",level:"3"},{id:"sec_10_3",title:"4.3.2 Good manufacturing practice",level:"3"},{id:"sec_13",title:"5. Conformity and other related concepts as characteristics of the ‘goods’",level:"1"},{id:"sec_13_2",title:"5.1 Brand",level:"2"},{id:"sec_15",title:"6. Burden of proof for the non-conformity of the goods with the standard",level:"1"},{id:"sec_15_2",title:"6.1 Burden of proof for the non-conformity of the goods at the time risk passes",level:"2"},{id:"sec_15_3",title:"6.1.1 Which risks are transferred to the buyer?",level:"3"},{id:"sec_15_4",title:"6.1.1.1 Liability for loss or damage to the goods",level:"4"},{id:"sec_19",title:"7. Proof as matter regulated under the uniform Act",level:"1"},{id:"sec_20",title:"8. Overview of the various allocation of burden of proof in practice",level:"1"},{id:"sec_21",title:"9. Conclusion",level:"1"}],chapterReferences:[{id:"B1",body:'\nPilkington N. The security of transactions and investment in Africa. In: Tumnde M, editor. Proceedings of the OHADA Seminar held at the University of Buea, Buea, Cameroon, 18-19 September 2003 on the Theme: The Applicability of the OHADA Treaty in Cameroon. Buea, Cameroon: University of Buea. pp. 28-41, pp. 28-30\n'},{id:"B2",body:'\nEbi NJ. Reflecting on OHADA law reform mission: Its impact on certain aspects of company law in Anglophone Cameroon. In: Annales de la Faculté des Sciences Juridiques et Politiques. Vol. 6. Université de Dschang; 2002. pp. 97-119 at 102\n'},{id:"B3",body:'\nSchlechtriem P. Requirements of application and sphere of applicability of the CISG. Victoria University of Wellington Law Review. 2005;35:781-794, 787\n'},{id:"B4",body:'\nCollin PH. Dictionary of Law. 3rd ed. Peter Collin Publishing; 2000. p. 77\n'},{id:"B5",body:'\nMba-Owono C. Non-conformité et Vices Caches dans la Vente Commerciale en Droit Uniform Africain [Non-conformity and hidden defects in commercial sales under the African Uniform Law]. Juridis Périodique. 2002;41:107, at 108\n'},{id:"B6",body:'\nSchwenzer I, editor. Commentary on the UN Convention on the International Sale of Goods (CISG). 3rd ed. Oxford University Press; 2010. p. 570\n'},{id:"B7",body:'\nFourment F. Défauts de la chose vendue que reste-t-il de l’action en garantie des vices cachés? [Hidden defects of sold goods: What action to take for hidden defects?]. Revue Trimestrielle de Droit Commercial et de Droit Economique. 1997;3:416, at 419\n'},{id:"B8",body:'\nAwona SPL. Défaut de conformité et défaut cache dans la vente commercial OHADA: Retour à la case départ? [Defects of conformity and hidden defects in the OHADA commercial sale: Coming back to the starting point?]. Recueil d’Études sur l’OHADA et les Normes Juridiques Africaines. 2013;VI:317-320\n'},{id:"B9",body:'\nPougoué PG, et al. Encyclopédie du Droit OHADA [Encyclopaedia of OHADA law] (2011, Lamy) at 55. See English Sale of Goods Act, secs 12-15 and CISG, arts 35-44\n'},{id:"B10",body:'\nHamilton WH. The ancient maxim caveat emptor. Yale Law Journal. 1928;40:1133 at 1186\n'},{id:"B11",body:'\nAriely D, Levav J. Sequential choice in group settings: Taking the road less traveled and less enjoyed. Journal of Consumer Research. 2000;27(3):279-290 at 279\n'},{id:"B12",body:'\nRatner RK, Kahn BE. The impact of private versus public consumption on variety-seeking behavior. Journal of Consumer Research. 2000;29(2):246-247\n'},{id:"B13",body:'\nLee JC, Kim J, Kwak K. A multi-attribute examination of consumer conformity in group-level ordering. Australasian Marketing Journal. 2018;26(1):41-48, at 41-42\n'},{id:"B14",body:'\nBridge M. The Sale of Goods. London: Oxford University Press. pp. 42-45\n'},{id:"B15",body:'\nSchlectreim PH. Non-material damages—Recovery under the CISG? Pace International Law Review. 2007;19:89-102, point III (3). Available from: http://www.cisg.law.pace.edu/cisg/biblio/schlechtriem15.html [Accessed: 07 April 2014]\n'},{id:"B16",body:'\nRatner RK, Kahn BE. The Impact of Private versus Public Consumption on Variety-Seeking Behavior., cit op. p. 246\n'},{id:"B17",body:'\nMudambi SMD, Doyle P, Wong V. An exploration of branding in industrial markets. Industrial Marketing Management. 1997;26(5):433-446, at 438\n'},{id:"B18",body:'\nMitchiner J. Intellectual property in image—A mere inconvenience. Intellectual Property Quarterly. 2003;2:163-208, at 182\n'},{id:"B19",body:'\nQuester P, Steyer A. Revisiting individual choices in group settings: The long and winding (less traveled) road? Journal of Consumer Research. 2010;36(6):1050-1057, at 1050-1051\n'},{id:"B20",body:'\nDjieufack R, Mbifi R, Nyiawung M. Quality Control of Goods and the Pricing Policy in Cameroon: A Legal Discourse. Germany: Galda Verlag; 2019. pp. 42-44\n'},{id:"B21",body:'\nHartman S. Brand equity impairment—The meaning of dilution. The Trademark Reporter. 1997;87:418-354 at 418\n'},{id:"B22",body:'\nSantos AP, Toé JY. OHADA Droit Commercial Général, Bruylant, collection Droit Uniforme Africain, Bruxelles; 2002. p. 405\n'},{id:"B23",body:'\nHenschel RF. Conformity of goods in international sales governed by CISG article 35: Caveat Venditor, Caveat Emptor and Contract Law as background law and as a comparing set of rules. Nordic Journal of Commercial Law:10\n'},{id:"B24",body:'\nSantos AP, Toé JY. OHADA Droit Commercial, above at note 34 at 392-393; Schwenzer (ed) Commentary, above at note 4 at 569\n'},{id:"B25",body:'\nStocks M. Risk of loss under the uniform commercial code and the convention on the international sale of goods: A comparative analysis and proposed revision of UCC sections 2-509 and 2-510. Northwestern University Law Review. 1993;87:1\n'},{id:"B26",body:'\nSaidov D. Article 35 of the CISG: Reflecting on the present and thinking about the future. Villanova Law Review;58\n'},{id:"B27",body:'\nRoberts T. Commercial impossibility and frustration of purpose: A critical analysis. Canadian Journal of Law & Jurisprudence. 2003;16:129, at 134\n'}],footnotes:[{id:"fn1",explanation:"The Treaty creating OHADA was signed at Port-Louis, Mauritius Island on 17 October 1993, as revised at Quebec, Canada, on 17 October 2008. The revisions became effective on 21 March 2010. As of July 7, 2010, the West African members of OHADA are Benin, Burkina Faso, Cote d’Ivoire, Guinea, Guinea-Bissau, Mali, Niger, Senegal, and Togo, and the Central African members of OHADA are Central African Republic, Chad, Cameroon, Comoros, Congo, Equatorial Guinea, and Gabon. See http:// and http://. On February 22, 2010, the Democratic Republic of Congo’s president ratified the country’s adoption of the OHADA treaty."},{id:"fn2",explanation:"The rules adopted are known as Uniform Acts."},{id:"fn3",explanation:"The following Uniform Acts are already applicable in Member States: Commercial Companies and Economic Interest Groupings, Law of Securities, Simplified Recovery Procedures and Measures of Execution, Collective Proceedings for Wiping –off Debts, Arbitration Law, Accounting Law, and Law of Co-operatives Carriage of Goods by Road. Two other Uniform Acts have been enacted and adopted by the Council of Ministers but are still inapplicable, to wit; Consumer Law and Contract Law."},{id:"fn4",explanation:"Article 30 UAGCL."},{id:"fn5",explanation:"Cameroon Civil Code, art 1641; Côte d’Ivoire Civil Code, art 1641."},{id:"fn6",explanation:"Uniform Act, art 231."},{id:"fn7",explanation:"SGA, sec 11(3) provides that a condition is a major term of a contract, breach of which is considered to go to the root of the contract so as to entitle the innocent party to treat the contract as discharged."},{id:"fn8",explanation:"Id, secs 14–15."},{id:"fn9",explanation:"CISG, art 35."},{id:"fn10",explanation:"Under French law, the hidden defect element is dealt with under sales law. This is actually effectively the purport of the text: Pougoué et al Encyclopedie, above at note 11 at 55."},{id:"fn11",explanation:"This is the author’s translation."},{id:"fn12",explanation:"The Moorcock (1889) 14 PD 64; Lister v Romford Ice Co Ltd [1957] AC 555."},{id:"fn13",explanation:'Compare with Lord Tomlin in Hillas & Co Ltd v Arcos [1932] ALL ER 494 at 499: "The problem for a court of construction must always be so to balance matters that, without violation of essential principle, the dealings of men may so far as possible be treated as effective, and that the law may not incur the reproach of being the destroyer of bargains."'},{id:"fn14",explanation:"Uniform Act, art 257."},{id:"fn15",explanation:"Id, art 259. Under CISG, art 39, the time frame is two years: J Huet, Contrats Civilset Commerciaux, Responsabilité du Vendeur et Garantie Contre les Vices Cachés [Civil and commercial contracts: Responsibility of the seller and guarantee against hidden defects] (1987, Litec) at 42."},{id:"fn16",explanation:"Uniform Act, art 255(2)."},{id:"fn17",explanation:"Sale of Goods Act § 14(2)–(2)(B)."},{id:"fn18",explanation:"As per the position in the civil codes of Cameroon (art 1641), Côte d’Ivoire (art 1641) and France (art 1641)."},{id:"fn19",explanation:"Id, art 283."},{id:"fn20",explanation:"Sale of Goods Act, 1979, c. 54, § 14(2)–(2)(B) (Eng.), available at http://www.legislation.gov.uk/ukpga/1979/54; U.C.C.§ 2-314(2)(c) (1977)."},{id:"fn21",explanation:"These would include the code of ethics of manufacturers’ associations, association of advertising practitioners, etc."},{id:"fn22",explanation:"Uniform Act, arts 238–239."},{id:"fn23",explanation:"Uniform Act, art 255 para. 1.0."},{id:"fn24",explanation:"Uniform Act, art 255 para. 2."},{id:"fn25",explanation:"Uniform Act, art 282-283."},{id:"fn26",explanation:"Uniform Act, arts 292-293."},{id:"fn27",explanation:"Definition of GMP by the World Health Organisation and as per GMP institute found at , Accessed on October 24th, 2009."},{id:"fn28",explanation:"It was created by Decree No.88/204 of 5 February 1988, re-organising the Ministry of Commerce. See Juridis Périodique no. 5 1991, p. 13. These structures were created with the objective of protecting consumers and ensuring the technical supervision of the quality and quantity of goods through inspection."},{id:"fn29",explanation:"Ibid, 438."},{id:"fn30",explanation:"Ibid, 438."},{id:"fn31",explanation:"Oxford English Dictionary (2nd edn, 1989), s.n. ‘brand’."},{id:"fn32",explanation:"Uniform Act, arts 256 & 258."},{id:"fn33",explanation:"Uniform Act, art 258."},{id:"fn34",explanation:"Uniform Act, art 256."},{id:"fn35",explanation:"Under art 1641 of the French Civil Code, for the purpose of an implied or legal guarantee, the defects must be hidden defects (vices cachés), unknown to the buyer."},{id:"fn36",explanation:"For example, in complex agreements, a part concerning services may entail a certain risk in conjunction with a sale but not governed by the Uniform Act. (Art 235(b))."},{id:"fn37",explanation:'This is the author’s translation of the French version of the UAGCL, which uses the words: "la perte ou la deterioration".'},{id:"fn38",explanation:"The Nkoulouloun and Mokolo markets respectively."},{id:"fn39",explanation:"Uniform Act, art 254."},{id:"fn40",explanation:"Uniform Act, art 278 para. 2."},{id:"fn41",explanation:"Uniform Act, art 281 & 296."},{id:"fn42",explanation:'"If the buyer rejects the goods by invoking their non-conformity the seller must prove that the goods are in conformity with the contract; if the buyer already accepted the goods the buyer would have to prove their non-conformity."'},{id:"fn43",explanation:"The author’s translation."},{id:"fn44",explanation:"Sometimes this rule is broken down into two separate rules distinguishing between the burden for a party raising a claim and a party claiming an exception or raising a defence."},{id:"fn45",explanation:"The author’s translation."}],contributors:[{corresp:"yes",contributorFullName:"Djieufack Roland",address:"orolly@yahoo.fr",affiliation:'
Higher Technical Teacher Training College, Bambili, The University of Bamenda, Bamenda, Cameroon
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Reporting",isOpenForSubmission:!1,hash:"05150c3d097ed7178a67213166db89b6",slug:"accounting-and-finance-new-perspectives-on-banking-financial-statements-and-reporting",bookSignature:"Reza Gharoie Ahangar and Can Öztürk",coverURL:"https://cdn.intechopen.com/books/images_new/8644.jpg",editedByType:"Edited by",editors:[{id:"91081",title:"Dr.",name:"Reza",surname:"Gharoie Ahangar",slug:"reza-gharoie-ahangar",fullName:"Reza Gharoie Ahangar"}],equalEditorOne:{id:"280253",title:"Dr.",name:"Can",middleName:null,surname:"Ozturk",slug:"can-ozturk",fullName:"Can Ozturk",profilePictureURL:"https://mts.intechopen.com/storage/users/280253/images/system/280253.jpg",biography:"Dr. Can Öztürk is an associate professor of financial, international and comparative accounting at the Department of Management, Çankaya University, Ankara, Turkey. He received his first Bachelor’s in Banking and Finance (2001) from Bilkent University, his second Bachelor\\'s in Finance (2003) and Master’s in Business Administration (2004) from the University of Massachusetts Dartmouth, USA as well as his Ph.D. in Accounting and Finance in 2011 from Başkent University, Turkey. His primary research interests are IFRS, IFRS for SMEs, Financial Reporting Standards for Non-Publicly Accountable Entities, ISAs, and Corporate Social Responsibility Reporting. He has published several articles in prestigious journals at the national and international context such as Accounting in Europe, Accounting and Management Information Systems, Emerging Markets Journal and The World of Accounting Science. 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Sison III",authors:[{id:"264901",title:"Mr.",name:"Maximo Paulino Iii",middleName:null,surname:"Sison",fullName:"Maximo Paulino Iii Sison",slug:"maximo-paulino-iii-sison"}]}]}]},onlineFirst:{chapter:{type:"chapter",id:"75844",title:"Pseudomonas aeruginosa Secreted Biomolecules and Their Diverse Functions in Biofilm Formation and Virulence",doi:"10.5772/intechopen.96866",slug:"pseudomonas-aeruginosa-secreted-biomolecules-and-their-diverse-functions-in-biofilm-formation-and-vi",body:'
1. Introduction
Pseudomonas aeruginosa is an opportunistic Gram-negative bacilli bacterium which holds a greater clinical significance in relation to its infection causing ability in humans [1]. P. aeruginosa is commonly found in environment (soil and water) and can be a source of contamination of drinking water and food spoilage [2, 3]. Prevalence of P. aeruginosa and its associated infection is commonly found in cystic fibrosis patient and chronic obstructive pulmonary disease (COPD) lungs, urinary tract, immunocompromised (HIV) patients, skin and soft-tissue, diabetic leg wounds, burns and surgical site infections [1, 4]. It is also a common pathogen responsible for causing healthcare associated (nosocomial) infection and microbial keratitis (eye infection due to contamination of contact lenses) [4]. World Health Organization (WHO) has listed P. aeruginosa as a most critical pathogen, due to the threat of causing blood stream infection (septicemia) and its antibiotic resistance ability [5]. P. aeruginosa or in general many other bacterial pathogens (e.g. Acinetobacter baumannii, Escherichia coli, Klebsiella pneumoniae, Staphylococcus aureus, Streptococcus epidermidis, Streptococcus pyogenes, Proteus mirabilis, etc.) has an inheritance ability to colonize and form biofilms on biotic (e.g. mucosa, tissue) or abiotic surface (e.g. medical implants, surgical instruments, hospital beds, wash basins, sinks, bath tub, etc.). Bacterial colonization on these surfaces directly leads to the contamination of surfaces, food and water and consequently precedes to infections in host. Bacterial biofilms are liable for approximately 80% of hospital and community-associated infections [6]. The most serious concern is antibiotic/antibacterial agents’ resistance by the infecting bacteria that threatens the very core of modern medicine and impose a greater burden on global public health and economy.
2. Bacterial infection and antibiotic resistance are a global concern
National Institutes of Health (NIH), USA statistics data reports 550,000 death a year and about $94 billion total cost annually associated with biofilm infections in USA alone [7]. In Australia, billions of dollars expended annually dealing with antibiotic-resistant infections [8]. Statistics on antibiotic-resistance bacteria causing healthcare associated infections (HAI’s) and death in European countries is alarming. Around, 8.9 million HAI recorded each year in combined hospitals and long-term care facilities and one in three bacteria associated with HAI’s are antibiotic resistant [9]. In European population death associated with antibiotic resistance bacteria is estimated to be around 33,000 annually, this statistic is comparable to death associated with combined influenza, tuberculosis, HIV/AIDS [10]. Antibiotic resistance associated infections also triggers massive loss in productivity and healthcare incidentals estimated to be approximately 1.5 billion Euro’s each year [11]. Bacterial resistance profile to different antibacterial agents is depends of several factors including geographical location of the strain (genetic mutations influenced by temperature, nutrient, oxygen), antibiotic prescribing practice to patients around the globe, poor hygiene and sanitation practice by common public and health care workers in hospitals. For instance, report released by Australian Commission on Safety and Quality in Health Care (Antimicrobial use and Resistance in Australia, AURA 2019) suggest more than 26.5 million antibiotic prescriptions were give out to patients [12]. The same report also highlighted that 23.5% of hospital prescribed antimicrobials in Australia are inappropriate and also community associated increase in antibiotic resistance bacteria (MRSA) are higher among old age people living in aged care facilities and in remote regions of the country [12]. Misuse or unethical use of antibiotics in agriculture, meat and poultry industry and fish farming, is a primary concern. Study published by Chinese Academy of Sciences reports use of 162,000 tons of antibiotics in the year 2013 alone in which more than half (52%) was used for animal husbandry and 48% by humans in addition, massive amount (50,000 tons) of antibiotics drained in the environment (water and soil) [13, 14]. USA also reported 10,000 tons of antibiotics annually used for livestock [15]. India, China, USA, Russia, Brazil, and South Africa are the world leaders in consuming antibiotics [16]. South China Morning Post (SCMP) Newspaper published an article in 2017 stating “Antibiotic overuse is a ticking time bomb for Asia” and health care workers act instantly to restrain misuse of antibiotics to stop public health calamity [17]. World Health Assembly, WHO, United Nations (UN), and countries respective government, local health organization and institutes are adopting a global action plan to crumb antibiotic resistance by educating common public, health care workers on effective sanitation, hygiene and infection prevention measures; and spreading awareness on responsible use of antibiotics in human and animal health, investing more funding in research and development in developing novel antibacterial agents, diagnostic tools, vaccines, improving hospital facilities especially in low-and-middle income countries [18].
3. P. aeruginosa Antibiotic resistance profile around the globe
P. aeruginosa inherent and adaptive antibiotic resistance character thus consequently making many existing antibiotics and anti-pseudomonal agents unusable against this bacterium and present a significant challenge for medical practitioners to treat infections. In this section, we exhibited few cases based on P. aeruginosa antibiotic resistance profile from different parts of the world by referring to previously published literature.
A comprehensive review by Wozniak et al., (2017), that covered the Australian data from the year 1990 till 2017, on antibiotic resistance Gram-negative bacteria [19]. Their study highlighted that P. aeruginosa isolates from different infection site showed resistance to many commonly used antibiotics. Among the P. aeruginosa isolates that were collected from surgical site between years 2002–2013, approximately 0.5%, 7.7% and 0.5% of the isolates showed resistance to fluoroquinolone, third generation cephalosporin and gentamicin respectively [20]. Survey on antibiotic resistance profile of P. aeruginosa isolates from patient’s sputum between years 2007–2010 showed resistance to aminoglycosides (43%), beta-lactam (21%) and fluoroquinolone (30%) class of antibiotics [21]. Epidemiology studies on P. aeruginosa isolates from blood (years: 2001–2009) showed resistance to fluoroquinolone and meropenem about 12.7% and 14.3% respectively [22]. National Healthcare Safety Network (NHSN), USA survey on antimicrobial resistance patterns for the year 2009–2010, reported about 20% of pathogens (from 69,475 HAI’s incidence) are antibiotic resistance in which 2% is carbapenem-resistant P. aeruginosa [23]. Microbial analysis on patients affected with Nosocomial and ventilator-associated pneumonia (VAP) in a period 2011–2012 in Georgia, USA reported P. aeruginosa as most prevalent Gram negative (40%) and highest prevalence of multi drug resistance [24]. Similar multi-drug resistance profile of P. aeruginosa was recorded in Asian countries. For example, antibiogram of total 2444 Pseudomonas species isolated from different clinical specimens (blood, pus, tracheal aspirate, urine and sputum from wards, intensive care units (ICUs) and follow up patients) of trauma patients from tertiary care hospitals in India over a period 2012–2016 revealed dominance of P. aeruginosa (95%) [25]. Among 69%, 68%, 67% 66%, 63% and 51% were levofloxacin, gentamicin, ciprofloxacin, ceftazidime, meropenem and tobramycin resistance, respectively [25]. Antibiotic profile of 121 P aeruginosa strains isolated from hospitals of Makkah and Jeddah, Saudi Arabia showed high resistance to antibiotics: meropenem (~30.6%), ticarcillin (22.3%), and imipenem (19%) [26]. A study reported that in mainland china hospitals prevalence of P. aeruginosa related ventilator-associated pneumonia (VAP) and hospital-acquired pneumonia were 19.4 and 17.8% respectively [27]. National Healthcare Safety Network (NHSN) USA, reports prevalence of P. aeruginosa is common among possible VAP [28]. These isolates exhibited high level of resistance to antibiotics: Gentamicin (up to 51.1%), cefoperazone (50%), and about 22.5% for amikacin [28]. P. aeruginosa resistance to ciprofloxacin has also risen a global concern, especially in Asian countries for example, Bangladesh reported 75.5% resistance to ciprofloxacin whereas, India, Iran, Turkey, and Saudi Arabia reported 49%, 58%, 48.9% and 50.9% respectively [29, 30, 31, 32, 33].
4. Role of P. aeruginosa secreted biomolecules in biofilm formation and virulence
Biofilm formation is the most preferred stage of many bacterial pathogens. Biofilm formation is a multi-step process to start with i) initial attachment of bacteria to the surface (adhesion) and to each other (aggregation), ii) growth regulations and microcolony formation and production of extracellular polymeric substances (EPS) and other exogenous molecules, iii) maturation of biofilms includes structural stability and iv) dispersal of bacterial cells from the mature biofilm into the environment and reestablishment at a new site [34].
Bacteria in its biofilm state are known to withstand antibacterial agents by many ten’s and 100’s-fold in comparison to its sessile/planktonic state [35]. Biofilm main composition includes up to 90% bacterially self-secreted biopolymers also known as extracellular polymeric substances (EPS) and other exogenous molecules and 10% bacterial cells [36]. These molecules in combined has been termed as house of bacteria and it shelter bacterial cells from numerous challenges includes antibiotics, antiseptics, detergents, shear mechanical stress, etc. [36]. Exogenous molecules synthesized by P. aeruginosa is primarily structured by a complex Quorum Sensing (QS) mechanism [37, 38]. In simple terms, QS is an intracellular communication phenomenon in which bacterial species able to detect and respond to its own cell population and ecological cues by regulating genes that facilitates them in survival and colonization in both biotic and abiotic environment. In P. aeruginosa QS is hierarchical and its driven through four known signaling system. At the top or first stage is driven by las system that activates the biosynthesis of autoinducing molecules N-(3-oxododecanoyl)-L-homoserine lactone (3-oxo-C12-HSL) and N-butanoyl-L-homoserine lactone (C4-HSL). Binding of LasR-HSL molecules triggers the transcription of second QS system: rhlR, rhlI, lasI. LasR system further regulates the third and fourth: 2-heptyl-3-hydroxy-4-quinolone (HHQ) and pseudomonas quinolone signal (PQS) [39]. These four QS circuits are interconnected and depends on each other regarding P. aeruginosa biosynthesis of various secreted and surface attached molecules. These includes extracellular biopolymers (Extracellular DNA, polysaccharides, proteins/enzymes), biosurfactant (rhamnolipids), metabolites (phenazine/pyocyanin), iron chelator (siderophore: pyoverdine, pyochelin), and bacterium cell surface anchored flagella and pili for swarming and twitching motilities [37, 38, 39, 40]. These biomolecules and cell appendages independently or in coordination with each other plays dominant role in P. aeruginosa growth, fitness, biofilm formation, virulence, pathogenicity in host during infection, antibacterial resistance, and persistence. In this chapter we emphasize only on the diverse role of P. aeruginosa secreted extracellular biomolecules. Figure 1 summarizes the diverse function of P. aeruginosa secreted extracellular biomolecules.
Figure 1.
Highlighting the major role of biomolecules secreted by P. aeruginosa. These biomolecules are essential for establishment of biofilm, bacterial growth, fitness, and survival, induce virulence/pathogenicity and triggering immune response in host during infection, evading antibiotics, and other antibacterial agents.
4.1 Extracellular DNA production, role in P. aeruginosa biofilm formation and stability
The role of extracellular (eDNA) in P. aeruginosa biofilm was first highlighted by Whitchurch et al. (2002) [41]. Their study revealed that eDNA is predominant in P. aeruginosa matrix component and its essential for P. aeruginosa biofilm formation [41]. Followed which numerous discoveries were done highlighting several roles of eDNA in P. aeruginosa and in other bacterial pathogens as well as in fungi [42, 43, 44, 45, 46]. Structural analysis study revealed that eDNA is similar to bacterial chromosomal DNA in its primary structure and it is not surprising because when chromosomal DNA release from bacterial cells (either via membrane vesicles or cell lysis) into its immediate environment is termed as eDNA [47]. eDNA in P. aeruginosa cell population is released primarily through QS mechanism [48]. QS system (las and rhl -acyl homoserine lactone and pqs-Pseudomonas quinolone signaling), as well as flagella and type IV pili (fliMpilA) facilities prophage induction in P. aeruginosa cell population and consequently trigger cell lysis and eDNA release [48]. Virulence factor pyocyanin/phenazine biosynthesis also shown to trigger cell lysis (via oxidative stress mediated by hydrogen peroxide) and eDNA release in P. aeruginosa [49]. Outer membrane vesicles in P. aeruginosa cell also demonstrated to actively release eDNA [50].
Studies have confirmed that eDNA plays a key role in different stages of biofilm formation including initial bacterial to surface attachment (adhesion), bacteria-to-bacteria interaction (aggregation), colonization and biofilm formation by connecting cells to cells like nanowires [41, 42, 43, 44, 45]. Presence of eDNA on P. aeruginosa cell surface have shown to dictates physical surface properties of bacterial cell such as increase in cell surface hydrophobicity and consequently enables physico-chemical interactions forces such as Van der Waals interactions, Acid–Base interactions, hydrophobic interactions that aids in bacterial interactions and biofilm formation [51, 52]. eDNA have proven to induce electrostatic interactions with divalent cations like calcium (Ca2+) and triggers bacterial aggregation [53]. eDNA has been established being an essential factor in structural integrity of P. aeruginosa biofilms and many studies have shown that cleaving of DNA using DNase I (enzyme that cleaves DNA through hydrolysis of phosphate di-ester bonds that links nucleotides in DNA) disrupts P. aeruginosa adhesion and biofilm formation [41, 44, 45, 54]. Other general roles of eDNA includes nutrient (e.g. good source of carbon, nitrogen, phosphorus) for starving bacteria and facilitate growth, horizontal gene transfer among bacteria cell (antibiotic resistance genes, virulence factor genes, etc), protects biofilms from shear stress by increasing biofilm viscosity. eDNA directly bindings to cationic antibiotics thus inhibits antimicrobial agents’ interaction with bacteria within biofilm, removal of eDNA from biofilms have shown increase of bacterial susceptibility to antimicrobial agents [55]. In P. aeruginosa biofilm, eDNA release has shown to lower the pH of the local environment and subsequently these acidification initiates antibiotic resistance phenotype genes (PhoPQ and PmrAB) that fosters alteration of lipid A and the manufacture of spermidine on the P. aeruginosa outer membrane and consequently decrease entry/intake of aminoglycoside antibiotics [56].
4.2 Multitude task of polysaccharides secreted by P. aeruginosa
Many studies have concluded that polysaccharides as a chief component of many bacterial EPS/biofilm matrix. P. aeruginosa biosynthesis alginate, psl, and pel as their three predominant extracellular polysaccharides. Alginate producing isolates of P. aeruginosa have been acknowledged as a mucoid phenotype regulates through mutation in the alginate biosynthesis of algA-algD operon and mucA [57]. AlgD is the key gene that promotes alginate production followed by combined action of mucA and algU genes [57]. The physical characterizes of alginate positive P. aeruginosa colonizes are highly viscous and gelatinous structure on the edge of the cells [58]. This feature is due to its heavy molecular weight structure of alginate which mainly composed of O-acetylated D-mannuronic acid and its C5′ epimer L-guluronic acid [59]. Alginate productions make P. aeruginosa virulent strain and a foremost cause for respiratory infections and mortality in CF patients [60]. Alginate production enhances bacterial adhesion due to its sticky nature and its plays key role in shielding P. aeruginosa from host immune defense system by scavenging reactive oxygen species (ROS) and evading neutrophils and macrophages mediated phagocytosis [61, 62]. A study by McCaslin in rat alveolar macrophages, showed that alginate in combination with lipopolysaccharide produced by P. aeruginosa plays a synergy role in sparking airway inflammation by impeding alveolar function in removal of apoptotic cells and debris [63]. The anionic (negative charge) feature of alginate undergoes electrostatic interactions with cationic aminoglycosides and thus constrains their dissemination into biofilms [64]. Alginate also induce structural and conformational alteration and aggregation in the antimicrobial peptides by binding with it thereby, hinders its antimicrobial activity against pseudomonas [65].
In absence of alginate biosynthesize, Psl or Pel genes in P. aeruginosa isolates up-regulates and activates over production of psl and pel polysaccharide [58]. These polysaccharides by itself or in combination with each other exhibit non-mucoid bacterial colonies/biofilm and these colonies are termed as rugose small colony variant (RSCV) [58]. Psl biosynthesis in P. aeruginosa is induced through a QS (las) mediated set of Psl genes (PslA-PslL) and each or group of Psl genes and its corresponding protein/enzyme plays a unique role in synthesizing and integrating Psl polysaccharide [58]. For instance, PslB enzyme is responsible for sugar-nucleotide precursor production, whereas, PslA/PslE/PslJ/PslK/PslL and PslF, PslH, and PslI set of enzymes deals with polymerization of polysaccharide, and integration of the activated sugar subunits into the polysaccharide repeating structure [58]. Psl is a neutrally charged polysaccharide comprised of repeating sugar groups: D-mannose, L-rhamnose, and D-glucose [66, 67]. This polysaccharide plays a crucial role in bacterial cell-to-cell communication by enhancing intracellular c-di-GMP (secondary messenger molecule) and essential for initial P. aeruginosa attachment to a surface as tested on various clinical, environmental and common laboratory strains, biofilm biomass and antibiotic tolerance (tested on gentamicin) [68, 69].
Pel is a positively charged polysaccharide comprised of amino sugar groups and is biosynthesized is regulated via QS (rhl sytem) through activating pel operons (pelA-pelG) [70, 71]. Pel composed of acetylated 1–4 glycosidic linkages of N-acetylgalactosamine and N-acetylglucosamine [71]. PelA protein is responsible for the deacetylase of the sugar amino group, whereas PelD, PelE, PelF, and PelG enzymes, these set of enzymes accountable for Pel polymerization and passage across the P. aeruginosa cytoplasmic membrane [58, 71]. Study also speculated that pel is adapted version of LPS [71]. Pel polysaccharide biosynthesize is a strain dependent, and studies shown that in absence of psl polysaccharides pel genes up regulated to form primary structural framework in non-mucoid P. aeruginosa biofilms. This indicates that pel plays important role on later stage of biofilm and not during initial adhesion, aggregation, and colonization [58]. Pel being a cationic biopolymer binds to negatively charged eDNA in P. aeruginosa biofilm matrix via ionic bonding/electrostatic interactions henceforth, stabilize biofilm matrix frame [72].
4.3 P. aeruginosa exotoxins proteins role in pathogenicity
The biosynthesize and secretion of exogenous proteins/enzymes by P. aeruginosa is mediated by QS (las-rhl) system [73]. The common proteins virulence factor P. aeruginosa secrets includes elastase/LAS A and B, exotoxin A, U, S, T, Y phospholipase C, alkaline protease, type IV protease, phospholipase H and lipolytic enzymes [74]. The primary function of these proteins is to play as a virulence factor and induce bacterial pathogenicity in host. To induce pathogenicity, evade host immune defense and damage epithelial cells, P. aeruginosa secrets these proteins predominantly via type II and type III secretion system (out of five protein secretion system) [75, 76]. Type II system constituent of protein secretons that facilitate release of exotoxin A, elastase/LasA and LasB proteases, type IV protease, and phospholipase H, as well as lipolytic enzymes into the host cells. Whereas exotoxins U, S, T, and Y are released into host cells via type III secretion system (T3SS) [76]. T3SS forms needle like membrane structure that are anchored to the bacterial cell surface and facilitates delivery of bacterial protein virulence factors into the host epithelial cells [76].
Some actions of P. aeruginosa virulence proteins are discussed below. For example, P. aeruginosa toxin A protein have shown to impair protein elongation factor in mammalian cells thereby interferes with host essential protein synthesis [77]. The T3SS proteins (Exo U, S, T, Y) have diverse functions such as hinder DNA synthesis and modulates cell morphology in host, escaping host phagocytosis by impairing host cell actin cytoskeleton polymerization and endothelial barriers, phospholipase activity (cleaving host cell lipid layer and increase cell membrane permeability), modulates host inflammatory response and consequently extending bacteria and its virulence factors into host blood stream, different organs to cause bacteraemia and septicaemia and organ failure [78, 79, 80]. Metalloproteases are another group of enzymes such as elastase whose main function is to cleave human elastin and leukocyte elastase and neutrophil elastase and consequently alters host tissue elastic property and stimulate tissue damage. Elastases also proven to degrade human collagen II and IV, impair fibroblast growth and destroy wound healing proteins which are essential for mammalian cell and tissue development and wound repair [81, 82, 83, 84]. Other crucial role of P. aeruginosa elastases includes cleaves host immunoglobulins (IgA and IgG) that aids bacterium to evade host immune response [85, 86]. Clinical studies in burn and wound patients infected with P. aeruginosa, showed protease biosynthesis by this bacterium trigger host cytokinin (interleukins IL6 and IL8) production and induce severe inflammation, septicaemia and elevates mortality level in patients [87, 88, 89].
4.4 Rhamnolipids P. aeruginosa biosurfactant
Rhamnolipids is a glycolipid biosurfactant produced by P. aeruginosa mediated through rhl QS system involving operons rhlA, rhlB for biosynthesis and rhlI and rhlR for regulation [90]. It is made up of sugar group (rhamnose) and a lipid/fatty acid group 3-(hydroxyalkanoyloxy) alkanoic acid and has a both hydrophilic and hydrophobic group like any typical biosurfactant [90, 91]. Rhamnolipids production helps P. aeruginosa in uptake and metabolism of hydrophobic molecules such as oils, hexadecane for nutritional source and growth [92]. Rhamnolipids (mono-rhamnolipids) also adhere to P. aeruginosa cell membrane (LPS) and plays key role in influencing P. aeruginosa cell surface physical property such as increasing cell surface hydrophobicity which aids in bacterial adhesion to substratum and bacterial cell-to-cell aggregation through hydrophobic interactions [93]. Rhamnolipids also lower the surface tension of P. aeruginosa cell surface thus aid them in swarming motility to travel across different location within the substratum [93]. It also proven to influence biofilm architecture by establishing and sustaining fluid channels in biofilms for water and oxygen transport [94]. P. aeruginosa employs rhamnolipids to their own advantage to eradicate competing bacteria. Binding of rhamnolipids into competing bacterial cell membrane consequently creates pores and increase cell permeability to induce cell lysis [95]. It is also a known virulence element, by binding to epithelial cell membrane it interrupts epithelial cell membrane integration, disrupts epithelial cell junctions, and triggers death in various mammalian cell types includes leukocytes, macrophages [96]. Rhamnolipids biosynthesis by P. aeruginosa in infected patients has been associated with escalation in pathogenicity in cystic fibrosis lung, ventilator-associated pneumonia patients [97].
4.5 Pyocyanin a unique virulence factor and its diverse function
P. aeruginosa biosynthesis and secretes a unique secondary metabolite called phenazines. Different types of phenazines are produced by P. aeruginosa however, pyocyanin is the most predominant one. Pyocyanin biosynthesis occurs at the later stage in P. aeruginosa population density or in biofilm, in laboratory culture it is generally expressed at the late exponential stage via regulation through QS (PQS) system [98]. Pyocyanin production is easily identified by its color, bluish -pure pyocyanin and green color when grown in laboratory in bacterial growth media (e.g. Tryptone Soy broth, Nutrient media, Luria broth, these media are all yellow in color and blue pyocyanin mix with yellow turns green). The two set genes of phzA1-G1 and phzA2-G2 encrypts initial phenazine molecule (phenazine-1-carboxylic acid, PCA) followed by conversion of PCA to pyocyanin (N-methyl-1-hydroxyphenazine) encoded by genes phzM and phzS [98]. Pyocyanin production has been associated with the severity of infection and acknowledged as a hyper virulent strain [99]. Analysis of pyocyanin production on variety of clinical and environmental isolates indicates pyocyanin production is very common among all isolates however, the amount of pyocyanin production is depended upon strain phenotype and genotype variations. A study by Fothergill et al. (2007) on strains isolated from different clinical sites (CF, keratitis) and environmental (water) strains indicated that Liverpool epidermic strain (LES) from CF patients (attended Liverpool CF centre in England between years 1995 to 2004) exhibited significantly high pyocyanin production in comparison to keratitis and water isolates [99]. Pyocyanin plays diverse role in establishment of P. aeruginosa biofilm formation including inducing oxidative stress in competing bacteria and outcompete their growth (e.g. S. aureus) and fungi (e.g. candida albicans) [100, 101]. Pyocyanin promote cell signaling by activating transcription factor SoxR and stimulating various genes expression includes efflux pump genes mexGHI-opmD, and PA2274 (monooxygenase, to control oxidative stress response in P. aeruginosa) [102]. By regulating target genes pyocyanin also maintain bacterial fitness, pyocyanin/phenazine deficient mutant (Δphz) showed drastic change in its colony morphology (wrinkled colony), whereas pyocyanin over producing mutant strain (DKN370) remained smooth [103]. Pyocyanin induce oxidative stress and cell death (via H2O2 production) in P. aeruginosa population in late exponential phase and triggers eDNA production [49]. An interesting discovery by Das et al. 2012 and 2015 revealed that pyocyanin intercalates with DNA and influence P. aeruginosa cell surface hydrophobicity and subsequently promote biofilm formation [51, 104].
Pyocyanin has been in limelight in many decades due to its virulence property. In context to P. aeruginosa infection in human, pyocyanin production has been linked to increase in virulence and severity of infection [99]. Different studies reported different concentration of pyocyanin to be found in sputum of CF patients from 0.9 to 16.5 μg/ml and 27.3 μg/ml in bronchitis patients sputum and also significantly higher amount (5.3 μg/g) also found in burn wound exudates [105, 106]. In mammalian cells, it declines intracellular cAMP and ATP levels, provoke neutrophils apoptosis, and modulates host immune system [105, 106, 107, 108]. Pyocyanin being a zwitter ion (positive and negative charge group and can penetrate into host cell membrane), and redox (electron donating and accepting property) molecule it oxidized cytosol (mammalian intracellular fluid), produces reactive oxygen species (ROS) by diffusing into host cells and undergoes redox reaction to accept electrons from NADPH and donates to molecular oxygen [109, 110]. ROS production leads to the decline in intracellular glutathione (a master antioxidant in mammalian cells essential for cell health and fitness) level which leads to bronchial epithelial cell death and tissue damage [109, 110]. It also impedes chlorine ion (Cl−) secretion and transport in CF patients’ lungs (bronchial epithelial cells) and consequently halt mucous clearance in human airways. [111]. In burn wound patients infected with P. aeruginosa, pyocyanin production shown to provoke premature senescence and apprehend human fibroblast growth by levying oxidative stress [106, 112]. Mouse model study revealed that exposing pyocyanin to mouse lung airways triggers repress of transcription factors protein FoxA2 expression (essential for tissue development) and consequently leads to over production of host cells (cell hyperplasia) and mucous hypersecretion by [113].
4.6 Siderophore benefits P. aeruginosa growth and biofilm formation
Siderophore are small molecules and belongs to the class of “iron-chelating compounds”. They are intrinsically secreted by microorganisms primarily for scavenging and uptake of Ferric ion, Fe3+ for their own benefits including nutrition, metabolism, growth, and virulence in mammals [114]. For example, Bacillus spp. (subtilis and anthracis) biosynthesis primary siderophore (bacillibactin), enterobactin, vibriobactin, yersinibactin, and pyoverdine by E.coli, Vibrio cholerae, Yersinia pestis and P. aeruginosa respectively. Pyoverdine is a fluorescent green color compound and its biosynthesis is encoded by the operons of pvd. Pyoverdine forages Fe3+ from host iron-binding molecules (transferrin) and binds strongly to it thus contribute to pathogenicity in host as shown in the immunocompromised mouse model [115, 116]. Pyoverdine also benefits from P. aeruginosa virulence factor protease action in degrading human iron-binding protein (ferritin), thus outcompetes host and scavenges iron [117]. Burn mouse model study have shown that pyoverdine contribute to severity in infection and mutants deficient in pyoverdine production showed significantly less virulence [116]. Infection model study in Caenorhabditis elegans, showed that pyoverdine penetrates host cells and undermines mitochondrial dynamics and triggers hypoxic response thus hinders ATP generation in host [118]. Other features of pyoverdine including communicating molecule to control biosynthesis of virulence proteins in P. aeruginosa including exotoxin A and protease [119]. Iron is essential to sustain bacterial growth thus pyoverdine aids in survival of P. aeruginosa in infection site, triggers biofilm formation where, pyoverdine deficient mutant strains forms fragile biofilm [120]. P. aeruginosa also produces another siderophore molecule called pyochelin, however pyochelin has lower affinity for Fe3+ than pyoverdine. However, this pyochelin-iron complex in coordination with pyocyanin undergoes oxidative-reductive reaction and contribute to oxidative damage (via hydroxyl radical formation) and inflammation in host [121, 122]. In CF patients pyochelin found to be involved in inflammation and tissue damage [123].
5. Conclusion
P. aeruginosa ability to easily colonize in host, biofilm formation, synthesis and secretion of virulence factors and causing pathogenicity, evading host immune defense system, and antimicrobial resistance made it a critical pathogen and needs an immediate attention. Secretion of extracellular molecules by P. aeruginosa plays a principal role in fitness of bacterial population, establishment of biofilms, infections, and pathogenicity in host. To reduce and eradicate P. aeruginosa associated infections development of novel antibiotics or antimicrobial agents, QS inhibiting molecules, virulence factor neutralizing agents, biofilm disrupting enzymes or/and combination treatment strategy with existing antimicrobial agents are of top priority. Further to prevent antimicrobial resistance in bacteria, necessary steps need to be taken by government organization, hospitals/clinics, health care workers and scientist from research institutes to educate children and students in schools, colleges and universities, people from rural places and developing countries about proper hygiene and use and misuse of antibiotics. Also, proper management of antibiotics uses in the agriculture and meat industry need to be implemented. This small steps at every level will help in minimize the spread of antimicrobial resistance in bacteria and will help to cut down catastrophe in both health and economic division and promotes better treatment outcome against infectious diseases.
\n',keywords:"Pseudomonas aeruginosa, pyocyanin, extracellular DNA, biofilms, alginate, rhamnolipids, pyoverdine",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/75844.pdf",chapterXML:"https://mts.intechopen.com/source/xml/75844.xml",downloadPdfUrl:"/chapter/pdf-download/75844",previewPdfUrl:"/chapter/pdf-preview/75844",totalDownloads:27,totalViews:0,totalCrossrefCites:0,dateSubmitted:"September 9th 2020",dateReviewed:"February 26th 2021",datePrePublished:"March 22nd 2021",datePublished:null,dateFinished:"March 22nd 2021",readingETA:"0",abstract:"Pseudomonas aeruginosa is an opportunistic Gram-negative bacterium accountable for causing life-threatening infections in humans. According to the World Health Organization, P. aeruginosa classified as a critical pathogen. Specifically, P. aeruginosa in its colonized or biofilm state presents a major infection threat to immunocompromised (HIV) patients, Cystic fibrosis, burns, wounds and surgery associated infection. It is also a common pathogen responsible for causing hospital acquired/nosocomial infection and Urinary tract infections. P. aeruginosa biofilm is made up of bacterial self-synthesized biomolecules includes extracellular DNA, polysaccharides, proteins, RNA, siderophores and metabolites such as pyocyanin. This chapter will elaborate the manifold functions of P. aeruginosa secreted biomolecules in establishing and stabilizing biofilms, triggering virulence and pathogenicity in host, and resisting antibiotics and antibacterial agents.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/75844",risUrl:"/chapter/ris/75844",signatures:"Theerthankar Das",book:{id:"9665",title:"Pseudomonas aeruginosa - Biofilm Formation, Infections and Treatments",subtitle:null,fullTitle:"Pseudomonas aeruginosa - Biofilm Formation, Infections and Treatments",slug:null,publishedDate:null,bookSignature:"Dr. Theerthankar Das",coverURL:"https://cdn.intechopen.com/books/images_new/9665.jpg",licenceType:"CC BY 3.0",editedByType:null,editors:[{id:"179493",title:"Dr.",name:"Theerthankar",middleName:null,surname:"Das",slug:"theerthankar-das",fullName:"Theerthankar Das"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:[{id:"179493",title:"Dr.",name:"Theerthankar",middleName:null,surname:"Das",fullName:"Theerthankar Das",slug:"theerthankar-das",email:"theerthankar_das11@rediffmail.com",position:null,institution:{name:"University of Sydney",institutionURL:null,country:{name:"Australia"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Bacterial infection and antibiotic resistance are a global concern",level:"1"},{id:"sec_3",title:"3. P. aeruginosa Antibiotic resistance profile around the globe",level:"1"},{id:"sec_4",title:"4. Role of P. aeruginosa secreted biomolecules in biofilm formation and virulence",level:"1"},{id:"sec_4_2",title:"4.1 Extracellular DNA production, role in P. aeruginosa biofilm formation and stability",level:"2"},{id:"sec_5_2",title:"4.2 Multitude task of polysaccharides secreted by P. aeruginosa",level:"2"},{id:"sec_6_2",title:"4.3 P. aeruginosa exotoxins proteins role in pathogenicity",level:"2"},{id:"sec_7_2",title:"4.4 Rhamnolipids P. aeruginosa biosurfactant",level:"2"},{id:"sec_8_2",title:"4.5 Pyocyanin a unique virulence factor and its diverse function",level:"2"},{id:"sec_9_2",title:"4.6 Siderophore benefits P. aeruginosa growth and biofilm formation",level:"2"},{id:"sec_11",title:"5. 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"Rhamnolipid surfactant production affects biofilm architecture in Pseudomonas aeruginosa PAO1. J Bacteriol. 2003;185(3):1027-36. doi:10.1128/jb.185.3.1027-1036.2003'},{id:"B95",body:'Sotirova AV, Spasova DI, Galabova DN, Karpenko E, Shulga A. Rhamnolipid-biosurfactant permeabilizing effects on gram-positive and gram-negative bacterial strains. Curr Microbiol. 2008;56 (6): 639-44. doi:10.1007/s00284-008-9139-3'},{id:"B96",body:'Zulianello L, Canard C, Köhler T, Caille D, Lacroix JS, Meda P. Rhamnolipids are virulence factors that promote early infiltration of primary human airway epithelia by Pseudomonas aeruginosa. Infect. Immun. 2006;74 (6): 3134-47. doi:10.1128/IAI.01772-05. PMC 1479292'},{id:"B97",body:'Köhler T, Guanella R, Carlet J, van Delden C. "Quorum sensing-dependent virulence during Pseudomonas aeruginosa colonisation and pneumonia in mechanically ventilated patients". Thorax. 2010;65 (8): 703-10. doi:10.1136/thx.2009.133082'},{id:"B98",body:'Mavrodi DV, Bonsall RF, Delaney SM, Soule MJ, Phillips G, Thomashow LS. Functional analysis of genes for biosynthesis of pyocyanin and phenazine-1-carboxamide from Pseudomonas aeruginosa PAO1. J Bacteriol. 2001;183(21):6454-6465'},{id:"B99",body:'Fothergill JL, Panagea S, Hart CA, Walshaw MJ, Tyrone L Pitt TL, Winstanley C. Widespread pyocyanin over-production among isolates of a cystic fibrosis epidemic strain. BMC Microbiol. 2007; 7: 45. Doi: 10.1186/1471-2180-7-45'},{id:"B100",body:'Sweden EG. Study the effect of antibiotics on pyocyanin production from Pseudomonas aeruginosa and pyocyanin as antibiotic against different pathogenic bacteria. Journal of University Anbar Pure Science. 2010; 4:15-18'},{id:"B101",body:'Kerr JR, Taylor GW, Rutman A, Hoiby N, Cole PJ, Wilson R. Pseudomonas aeruginosa pyocyanin and 1-hydroxyphenazine inhibit fungal growth. Journal of Clinical Pathology. 1999; 52:385-387'},{id:"B102",body:'Dietrich LEP, Price-Whelan A, Petersen A, Whiteley M, Newman DK. The phenazine pyocyanin is a terminal signalling factor in the quorum sensing network of Pseudomonas aeruginosa. 2006;61(5):1308-1321'},{id:"B103",body:'Dietrich LEP, Teal TK, Price-Whelan A, Newman DK Redox-Active Antibiotics Control Gene Expression and Community Behavior in Divergent Bacteria. Science. 2008; 321(5893): 1203-1206'},{id:"B104",body:'Das T, Kutty SK, Tavallaie R, Ibugo AI, Panchompoo J, Sehar S, Aldous L, Yeung AWS, Thomas SR, Kumar N, Gooding JJ, Manefield M. Phenazine virulence factor binding to extracellular DNA is important for Pseudomonas aeruginosa biofilm formation. 2015;5(8398)'},{id:"B105",body:'Wilson R, Sykes DA, Watson D, Rutman A, Taylor GW, Cole PJ. Measurement of Pseudomonas aeruginosa phenazine pigments in sputum and assessment of their contribution to sputum sol toxicity for respiratory epithelium. Infection and Immunity. 1988;56(9):2515-2517'},{id:"B106",body:'Muller M, Li Z, Maitz PK. Pseudomonas pyocyanin inhibits wound repair by inducing premature cellular senescence: role for p38 mitogen-activated protein kinase. Burns. 2009;35(4):500-508'},{id:"B107",body:'Munro NC, Barker A, Rutman A, Taylor G, Watson D, McDonald-Gibson WJ, Towart R, Taylor WA, Wilson R, Cole PJ. Effect of pyocyanin and 1-hydroxyphenazine on in vivo tracheal mucus velocity. J Appl Physiol (1985). 1989;67(1):316-323'},{id:"B108",body:'Denning GM, Wollenweber LA, Railsback MA, Cox CD, Stoll LL, Britigan BE. Pseudomonas pyocyanin increases interleukin-8 expression by human airway epithelial cells. Infect Immun. 1998;66(12):5777-5784'},{id:"B109",body:'O\'Malley YQ, Reszka KJ, Spitz DR, Denning GM, Britigan BE. Pseudomonas aeruginosa pyocyanin directly oxidizes glutathione and decreases its levels in airway epithelial cells. Am J Physiol Lung Cell Mol Physiol. 2004;287(1):L94-103'},{id:"B110",body:'Rada B, Jendrysik MA, Pang L, Hayes CP, Yoo DG, Park JJ, Moskowitz SM, Malech HL, Leto TL. Pyocyanin-enhanced neutrophil extracellular trap formation requires the NADPH oxidase. PLoS One. 2013;8(1):e54205'},{id:"B111",body:'Schwarzer C, Fischer H, Kim EJ, Barber KJ, Mills AD, et al. Oxidative Stress By Pyocyanin Impairs CFTR Cl− Transport In Human Bronchial Epithelial Cells. Free Radic Biol Med. 2008; 45(12): 1653-1662. Doi: 10.1016/j.freeradbiomed.2008.09.011'},{id:"B112",body:'Gonzalez MR, Fleuchot B, Lauciello L, Jafari P, Applegate LA, Raffoul W, Que YA, Perron K. Effect of human burn wound exudate on Pseudomonas aeruginosa virulence. mSphere. 2016;1(2)'},{id:"B113",body:'Hao Y, et al. Pseudomonas aeruginosa pyocyanin causes airway goblet cell hyperplasia and metaplasia and mucus hypersecretion by inactivating the transcriptional factor FoxA2. Cellular microbiology. 2012;14:401-415]'},{id:"B114",body:'Challis GL (April 2005). "A widely distributed bacterial pathway for siderophore biosynthesis independent of nonribosomal peptide synthetases". ChemBioChem. 6 (4): 601-11. doi:10.1002/cbic.200400283. PMID 15719346'},{id:"B115",body:'Sriyosachati S, Cox CD. Siderophore-mediated iron acquisition from transferrin by Pseudomonas aeruginosa. Infect Immun. 1986;52(3):885-891'},{id:"B116",body:'Meyer JM, Neely A, Stintzi A, Georges C, Holder IA. Pyoverdin is essential for virulence of Pseudomonas aeruginosa. Infection and Immunity. 1996;64(2):518-523'},{id:"B117",body:'Doring G, Pfestorf M, Botzenhart K, Abdallah MA. Impact of proteases on iron uptake of Pseudomonas aeruginosa pyoverdin from transferrin and lactoferrin. Infect Immun. 1988;56(1):291-293'},{id:"B118",body:'Kirienko, Natalia V.; Kirienko, Daniel R.; Larkins-Ford, Jonah; Wählby, Carolina; Ruvkun, Gary; Ausubel, Frederick M. (2013-04-17). "Pseudomonas aeruginosa disrupts Caenorhabditis elegans iron homeostasis, causing a hypoxic response and death". Cell Host & Microbe. 13 (4): 406-416. doi:10.1016/j.chom.2013.03.00'},{id:"B119",body:'Lamont IL, Beare PA, Ochsner U, Vasil AI, Vasil ML. Siderophore-mediated signaling regulates virulence factor production in Pseudomonas aeruginosa. Proc Natl Acad Sci U S A. 2002;99(10):7072-7077'},{id:"B120",body:'Lamont IL, Konings AF, Reid DW. Iron acquisition by Pseudomonas aeruginosa in the lungs of patients with cystic fibrosis. Biometals. 2009;22(1):53-60'},{id:"B121",body:'Coffman TJ, Cox, CD, Edeker BL., and Britigan BE. Possible role of bacterial siderophores in inflammation. Iron bound to the Pseudomonas siderophore pyochelin can function as a hydroxyl radical catalyst. J. Clin. Invest. 1990;86, 1030-1037. doi: 10.1172/JCI114805'},{id:"B122",body:'Britigan BE, Rasmussen G., Cox CD. Augmentation of oxidant injury to human pulmonary epithelial cells by the Pseudomonas aeruginosa siderophore pyochelin. Infect. Immun. 1997;65, 1071-1076'},{id:"B123",body:'Lyczak JB, Cannon CL,Pier GB. Lung infections associated with cystic fibrosis. Clin Microbiol Rev. 2002;15, 194-222. doi: 10.1128/CMR.15.2.194-222.2002'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Theerthankar Das",address:"das.ashishkumar@sydney.edu.au",affiliation:'
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BKCI is a part of Web of Science Core Collection (WoSCC) and the world’s leading citation index with multidisciplinary content from the top tier international and regional journals, conference proceedings, and books. The Book Citation Index includes over 104,500 editorially selected books, with 10,000 new books added each year. Containing more than 53.2 million cited references, coverage dates back from 2005 to present. The Book Citation Index is multidisciplinary, covering disciplines across the sciences, social sciences, and arts & humanities.
Produced by the Web Of Science group, BIOSIS Previews research database provides researchers with the most current sources of life sciences information, including journals, conferences, patents, books, review articles, and more. Researchers can also access multidisciplinary coverage via specialized indexing such as MeSH disease terms, CAS registry numbers, Sequence Databank Numbers and Major Concepts.
Produced by the Web Of Science group, Zoological Record is the world’s oldest continuing database of animal biology. It is considered the world’s leading taxonomic reference, and with coverage back to 1864, has long acted as the world’s unofficial register of animal names. The broad scope of coverage ranges from biodiversity and the environment to taxonomy and veterinary sciences.
Provides a simple way to search broadly for scholarly literature. Includes peer-reviewed papers, theses, books, abstracts and articles, from academic publishers, professsional societies, preprint repositories, universities and other scholarly organizations. Google Scholar sorts articles by weighing the full text of each article, the author, the publication in which the article appears, and how often the article has been cited in other scholarly literature, so that the most relevant results are returned on the first page.
Microsoft Academic is a project exploring how to assist human conducting scientific research by leveraging machine’s cognitive power in memory, computation, sensing, attention, and endurance. Re-launched in 2016, the tool features an entirely new data structure and search engine using semantic search technologies. The Academic Knowledge API offers information retrieval from the underlying database using REST endpoints for advanced research purposes.
The national library of the United Kingdom includes 150 million manuscripts, maps, newspapers, magazines, prints and drawings, music scores, and patents. Online catalogues, information and exhibitions can be found on its website. The library operates the world's largest document delivery service, providing millions of items a year to national and international customers.
The digital NSK portal is the central gathering place for the digital collections of the National and University Library (NSK) in Croatia. It was established in 2016 to provide access to the Library’s digital and digitized material collections regardless of storage location. The digital NSK portal enables a unified search of digitized material from the NSK Special Collections - books, visual material, maps and music material. From the end of 2019, all thematic portals are available independently: Digital Books, Digitized Manuscripts, Digitized Visual Materials, Digital Music Materials and Digitized Cartographic Materials (established in 2017). Currently available only in Croatian.
The official DOI (digital object identifier) link registration agency for scholarly and professional publications. Crossref operates a cross-publisher citation linking system that allows a researcher to click on a reference citation on one publisher’s platform and link directly to the cited content on another publisher’s platform, subject to the target publisher’s access control practices. This citation-linking network covers millions of articles and other content items from several hundred scholarly and professional publishers.
Dimensions is a next-generation linked research information system that makes it easier to find and access the most relevant information, analyze the academic and broader outcomes of research, and gather insights to inform future strategy. Dimensions delivers an array of search and discovery, analytical, and research management tools, all in a single platform. Developed in collaboration with over 100 leading research organizations around the world, it brings together over 128 million publications, grants, policy, data and metrics for the first time, enabling users to explore over 4 billion connections between them.
The primary aim of DOAB (Directory of Open Access Books) is to increase discoverability of Open Access books. Metadata will be harvestable in order to maximize dissemination, visibility and impact. Aggregators can integrate the records in their commercial services and libraries can integrate the directory into their online catalogues, helping scholars and students to discover the books.
OAPEN is dedicated to open access, peer-reviewed books. OAPEN operates two platforms, the OAPEN Library (www.oapen.org), a central repository for hosting and disseminating OA books, and the Directory of Open Access Books (DOAB, www.doabooks.org), a discovery service for OA books.
OpenAIRE aims at promoting and implementing the directives of the European Commission (EC) and the European Research Council on the promotion and funding of science and research. OpenAIRE supports the Open Access Mandate and the Open Research Data Pilot developed as part of the Horizon 2020 projects.
An integrated information service combining reference databases, subscription management, online journals, books and linking services. Widely used by libraries, schools, government institutions, medical institutions, corporations and others.
SFX® link resolver gives patrons and librarians a wealth of features that optimize management of and access to resources. It provides patrons with a direct route to electronic full-text records through OpenURL linking, delivers alternative links for further resource discovery, access to journals, and more. Released in 2001 as the first OpenURL resolver, SFX is continuously enhanced to support the newest industry developments and meet the evolving needs of customers. The records include a mix of scholarly material – primarily articles and e-books – but also conference proceedings, newspaper articles, and more.
A non-profit, membership, computer library service and research organization dedicated to the public purposes of furthering access to the world's information and reducing information costs. More than 41,555 libraries in 112 countries and territories around the world use OCLC services to locate, acquire, catalogue, lend and preserve library materials.
The world’s largest collection of open access research papers. CORE's mission is to aggregate all open access research outputs from repositories and journals worldwide and make them available to the public. In this way CORE facilitates free unrestricted access to research for all.
Perlego is a digital online library focusing on the delivery of academic, professional and non-fiction eBooks. It is a subscription-based service that offers users unlimited access to these texts for the duration of their subscription, however IntechOpen content integrated on the platform will always be available for free. They have been billed as “the Spotify for Textbooks” by the Evening Standard. Perlego is based in London but is available to users worldwide.
MyScienceWork provides a suite of data-driven solutions for research institutions, scientific publishers and private-sector R&D companies. MyScienceWork's comprehensive database includes more than 90 million scientific publications and 12 million patents.
CNKI (China National Knowledge Infrastructure) is a key national information construction project under the lead of Tsinghua University, and supported by PRC Ministry of Education, PRC Ministry of Science, Propaganda Department of the Communist Party of China and PRC General Administration of Press and Publication. CNKI has built a comprehensive China Integrated Knowledge Resources System, including journals, doctoral dissertations, masters' theses, proceedings, newspapers, yearbooks, statistical yearbooks, ebooks, patents, standards and so on. CNKI keeps integrating new contents and developing new products in 2 aspects: full-text academic resources, software on digitization and knowledge management. Began with academic journals, CNKI has become the largest and mostly-used academic online library in China.
As one of the largest digital content platform in China,independently developed by CNPIEC, CNPeReading positions herself as “One Platform,Vast Content, Global Services”. Through their new cooperation model and service philosophy, CNPeReading provides integrated promotion and marketing solutionsfor upstream publishers, one-stop, triune, recommendation, online reading and management servicesfor downstream institutions & libraries.
ERIC (Education Resources Information Center), sponsored by the Institute of Education Sciences (IES) of the U.S. Department of Education, provides access to education literature to support the use of educational research and information to improve practice in learning, teaching, educational decision-making, and research. The ERIC website is available to the public for searching more than one million citations going back to 1966.
The ACM Digital Library is a research, discovery and networking platform containing: The Full-Text Collection of all ACM publications, including journals, conference proceedings, technical magazines, newsletters and books. A collection of curated and hosted full-text publications from select publishers.
BASE (Bielefeld Academic Search Engine) is one of the world's most voluminous search sengines especially for academic web resources, e.g. journal articles, preprints, digital collections, images / videos or research data. BASE facilitates effective and targeted searches and retrieves high quality, academically relevant results. Other than search engines like Google or Bing BASE searches the deep web as well. The sources which are included in BASE are intellectually selected (by people from the BASE team) and reviewed. That's why data garbage and spam do not occur.
Zentralblatt MATH (zbMATH) is the world’s most comprehensive and longest-running abstracting and reviewing service in pure and applied mathematics. It is edited by the European Mathematical Society (EMS), the Heidelberg Academy of Sciences and Humanities and FIZ Karlsruhe. zbMATH provides easy access to bibliographic data, reviews and abstracts from all areas of pure mathematics as well as applications, in particular to natural sciences, computer science, economics and engineering. It also covers history and philosophy of mathematics and university education. All entries are classified according to the Mathematics Subject Classification Scheme (MSC 2020) and are equipped with keywords in order to characterize their particular content.
IDEAS is the largest bibliographic database dedicated to Economics and available freely on the Internet. Based on RePEc, it indexes over 3,100,000 items of research, including over 2,900,000 that can be downloaded in full text. RePEc (Research Papers in Economics) is a large volunteer effort to enhance the free dissemination of research in Economics which includes bibliographic metadata from over 2,000 participating archives, including all the major publishers and research outlets. IDEAS is just one of several services that use RePEc data.
As the authoritative source for chemical names, structures and CAS Registry Numbers®, the CAS substance collection, CAS REGISTRY®, serves as a universal standard for chemists worldwide. Covering advances in chemistry and related sciences over the last 150 years, the CAS content collection empowers researchers, business leaders, and information professionals around the world with immediate access to the reliable information they need to fuel innovation.
BKCI is a part of Web of Science Core Collection (WoSCC) and the world’s leading citation index with multidisciplinary content from the top tier international and regional journals, conference proceedings, and books. The Book Citation Index includes over 104,500 editorially selected books, with 10,000 new books added each year. Containing more than 53.2 million cited references, coverage dates back from 2005 to present. The Book Citation Index is multidisciplinary, covering disciplines across the sciences, social sciences, and arts & humanities.
Produced by the Web Of Science group, BIOSIS Previews research database provides researchers with the most current sources of life sciences information, including journals, conferences, patents, books, review articles, and more. Researchers can also access multidisciplinary coverage via specialized indexing such as MeSH disease terms, CAS registry numbers, Sequence Databank Numbers and Major Concepts.
Produced by the Web Of Science group, Zoological Record is the world’s oldest continuing database of animal biology. It is considered the world’s leading taxonomic reference, and with coverage back to 1864, has long acted as the world’s unofficial register of animal names. The broad scope of coverage ranges from biodiversity and the environment to taxonomy and veterinary sciences.
Provides a simple way to search broadly for scholarly literature. Includes peer-reviewed papers, theses, books, abstracts and articles, from academic publishers, professsional societies, preprint repositories, universities and other scholarly organizations. Google Scholar sorts articles by weighing the full text of each article, the author, the publication in which the article appears, and how often the article has been cited in other scholarly literature, so that the most relevant results are returned on the first page.
Microsoft Academic is a project exploring how to assist human conducting scientific research by leveraging machine’s cognitive power in memory, computation, sensing, attention, and endurance. Re-launched in 2016, the tool features an entirely new data structure and search engine using semantic search technologies. The Academic Knowledge API offers information retrieval from the underlying database using REST endpoints for advanced research purposes.
The national library of the United Kingdom includes 150 million manuscripts, maps, newspapers, magazines, prints and drawings, music scores, and patents. Online catalogues, information and exhibitions can be found on its website. The library operates the world's largest document delivery service, providing millions of items a year to national and international customers.
The digital NSK portal is the central gathering place for the digital collections of the National and University Library (NSK) in Croatia. It was established in 2016 to provide access to the Library’s digital and digitized material collections regardless of storage location. The digital NSK portal enables a unified search of digitized material from the NSK Special Collections - books, visual material, maps and music material. From the end of 2019, all thematic portals are available independently: Digital Books, Digitized Manuscripts, Digitized Visual Materials, Digital Music Materials and Digitized Cartographic Materials (established in 2017). Currently available only in Croatian.
The official DOI (digital object identifier) link registration agency for scholarly and professional publications. Crossref operates a cross-publisher citation linking system that allows a researcher to click on a reference citation on one publisher’s platform and link directly to the cited content on another publisher’s platform, subject to the target publisher’s access control practices. This citation-linking network covers millions of articles and other content items from several hundred scholarly and professional publishers.
Dimensions is a next-generation linked research information system that makes it easier to find and access the most relevant information, analyze the academic and broader outcomes of research, and gather insights to inform future strategy. Dimensions delivers an array of search and discovery, analytical, and research management tools, all in a single platform. Developed in collaboration with over 100 leading research organizations around the world, it brings together over 128 million publications, grants, policy, data and metrics for the first time, enabling users to explore over 4 billion connections between them.
The primary aim of DOAB (Directory of Open Access Books) is to increase discoverability of Open Access books. Metadata will be harvestable in order to maximize dissemination, visibility and impact. Aggregators can integrate the records in their commercial services and libraries can integrate the directory into their online catalogues, helping scholars and students to discover the books.
OAPEN is dedicated to open access, peer-reviewed books. OAPEN operates two platforms, the OAPEN Library (www.oapen.org), a central repository for hosting and disseminating OA books, and the Directory of Open Access Books (DOAB, www.doabooks.org), a discovery service for OA books.
OpenAIRE aims at promoting and implementing the directives of the European Commission (EC) and the European Research Council on the promotion and funding of science and research. OpenAIRE supports the Open Access Mandate and the Open Research Data Pilot developed as part of the Horizon 2020 projects.
An integrated information service combining reference databases, subscription management, online journals, books and linking services. Widely used by libraries, schools, government institutions, medical institutions, corporations and others.
SFX® link resolver gives patrons and librarians a wealth of features that optimize management of and access to resources. It provides patrons with a direct route to electronic full-text records through OpenURL linking, delivers alternative links for further resource discovery, access to journals, and more. Released in 2001 as the first OpenURL resolver, SFX is continuously enhanced to support the newest industry developments and meet the evolving needs of customers. The records include a mix of scholarly material – primarily articles and e-books – but also conference proceedings, newspaper articles, and more.
A non-profit, membership, computer library service and research organization dedicated to the public purposes of furthering access to the world's information and reducing information costs. More than 41,555 libraries in 112 countries and territories around the world use OCLC services to locate, acquire, catalogue, lend and preserve library materials.
The world’s largest collection of open access research papers. CORE's mission is to aggregate all open access research outputs from repositories and journals worldwide and make them available to the public. In this way CORE facilitates free unrestricted access to research for all.
Perlego is a digital online library focusing on the delivery of academic, professional and non-fiction eBooks. It is a subscription-based service that offers users unlimited access to these texts for the duration of their subscription, however IntechOpen content integrated on the platform will always be available for free. They have been billed as “the Spotify for Textbooks” by the Evening Standard. Perlego is based in London but is available to users worldwide.
MyScienceWork provides a suite of data-driven solutions for research institutions, scientific publishers and private-sector R&D companies. MyScienceWork's comprehensive database includes more than 90 million scientific publications and 12 million patents.
CNKI (China National Knowledge Infrastructure) is a key national information construction project under the lead of Tsinghua University, and supported by PRC Ministry of Education, PRC Ministry of Science, Propaganda Department of the Communist Party of China and PRC General Administration of Press and Publication. CNKI has built a comprehensive China Integrated Knowledge Resources System, including journals, doctoral dissertations, masters' theses, proceedings, newspapers, yearbooks, statistical yearbooks, ebooks, patents, standards and so on. CNKI keeps integrating new contents and developing new products in 2 aspects: full-text academic resources, software on digitization and knowledge management. Began with academic journals, CNKI has become the largest and mostly-used academic online library in China.
As one of the largest digital content platform in China,independently developed by CNPIEC, CNPeReading positions herself as “One Platform,Vast Content, Global Services”. Through their new cooperation model and service philosophy, CNPeReading provides integrated promotion and marketing solutionsfor upstream publishers, one-stop, triune, recommendation, online reading and management servicesfor downstream institutions & libraries.
ERIC (Education Resources Information Center), sponsored by the Institute of Education Sciences (IES) of the U.S. Department of Education, provides access to education literature to support the use of educational research and information to improve practice in learning, teaching, educational decision-making, and research. The ERIC website is available to the public for searching more than one million citations going back to 1966.
The ACM Digital Library is a research, discovery and networking platform containing: The Full-Text Collection of all ACM publications, including journals, conference proceedings, technical magazines, newsletters and books. A collection of curated and hosted full-text publications from select publishers.
BASE (Bielefeld Academic Search Engine) is one of the world's most voluminous search sengines especially for academic web resources, e.g. journal articles, preprints, digital collections, images / videos or research data. BASE facilitates effective and targeted searches and retrieves high quality, academically relevant results. Other than search engines like Google or Bing BASE searches the deep web as well. The sources which are included in BASE are intellectually selected (by people from the BASE team) and reviewed. That's why data garbage and spam do not occur.
Zentralblatt MATH (zbMATH) is the world’s most comprehensive and longest-running abstracting and reviewing service in pure and applied mathematics. It is edited by the European Mathematical Society (EMS), the Heidelberg Academy of Sciences and Humanities and FIZ Karlsruhe. zbMATH provides easy access to bibliographic data, reviews and abstracts from all areas of pure mathematics as well as applications, in particular to natural sciences, computer science, economics and engineering. It also covers history and philosophy of mathematics and university education. All entries are classified according to the Mathematics Subject Classification Scheme (MSC 2020) and are equipped with keywords in order to characterize their particular content.
IDEAS is the largest bibliographic database dedicated to Economics and available freely on the Internet. Based on RePEc, it indexes over 3,100,000 items of research, including over 2,900,000 that can be downloaded in full text. RePEc (Research Papers in Economics) is a large volunteer effort to enhance the free dissemination of research in Economics which includes bibliographic metadata from over 2,000 participating archives, including all the major publishers and research outlets. IDEAS is just one of several services that use RePEc data.
As the authoritative source for chemical names, structures and CAS Registry Numbers®, the CAS substance collection, CAS REGISTRY®, serves as a universal standard for chemists worldwide. Covering advances in chemistry and related sciences over the last 150 years, the CAS content collection empowers researchers, business leaders, and information professionals around the world with immediate access to the reliable information they need to fuel innovation.
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