Biological control of citrus postharvest phytopathogens: History Edit

Citrus is one of the most produced fruit genus. Grown in more than 100 countries, this group is composed by several species, including oranges, tangerines, mandarins, grapefruits, lemons, and limes [1]. The impact of citrus agroindustry in the international economy is huge. Besides their value as commodities, they also provide employment in many segments involved in its production cycle: harvesting, handling, transportation, and storage. In 2017, the global orange production reached 47.6 million metric tons (tons) and is expected to expand 4.2 million in 2018/2019 due to favorable weather in Brazil and United States, two of the most important orange producers in the world [2].

Over 20 different kinds of postharvest diseases have been reported in citrus and they are the main cause of fruit spoilage, resulting in massive economic losses [3]. Moreover, fungal growth in fruit may lead to production of mycotoxins, including potential carcinogenic agents such as citrinin and patulin [4], as well as tremorgenic compounds, for example tryptoquivalines [5], therefore representing a threat to human and animal life.

Green mold, blue mold, and sour rot, caused by Penicillium digitatum, P. italicum, and Geotrichum citri-aurantii, respectively, are the main citrus postharvest diseases [6]. P. digitatum, alone, is responsible for approximately 90% of total postharvest losses [7]. The fruits are contaminated through skin postharvest damage during their picking, packaging, storage, and transportation [3].

The interaction between citrus fruit and these phytopathogens is not fully understood, but some factors are known to affect this interaction in order to increase the fungus pathogenicity. P. digitatum and P. italicum are known to secrete organic acids during infection, leading to an optimal pH for its cell wall-degrading enzymes, such as polygalacturonases (PG) [8,9]. Moreover, P. digitatum also produce catalase during infection, an antioxidant enzyme that decomposes hydrogen peroxide, the main defense mechanism in citrus [10].

In order to deal with these fungi, chemical fungicides have been the main focus of research over the past decades. P. digitatum and P. italicum can be efficiently controlled by imazalil, thiabendazole, or pyrimethanil, but these fungicides are not effective against sour rot. For the control of G. citri-aurantii, guazatine, and propiconazole can be applied, although they are not allowed in some producing countries such as Brazil [11]. However, widespread use of chemical fungicides has caused the proliferation of resistant strains of these phytophatogens, compromising the effectiveness of these treatments [12]. Furthermore, concerns about environmental contamination and risks associated to human health have been raised around the accumulation of their residues in food.

Finding commercially viable, effective, alternative control methods has been a leading challenge for researchers, especially for controlling G. citri-aurantii, since there are fewer options available of acceptable chemical fungicides. Many alternatives have been proposed, including the application of antagonistic microorganisms and natural antimicrobial substances. Natural antimicrobial substances, especially plant extracts, are considered relatively safe, presenting low toxicity and high decomposability due to their natural origin, raising particular interest for use in these natural products [13].

Thus, the application of biocontrol agents has been an alternative for synthetic chemical fungicides. However, more research is necessary to understand their mechanism of action and effectiveness in different infection levels; this knowledge is crucial to implement their use as practical control agents. Therefore, the use of alternative postharvest biological control methods, both non-polluting and possessing low toxicity are reviewed here, highlighting advances presented in the literature in the recent years.