Contents lists available at ScienceDirect Journal of Biotechnology journal homepage: www.elsevier.com/locate/jbiotec Review Nepenthes: State of the art of an inspiring plant for biotechnologists Sissi Miguel a , Alain Hehn b,c, ⁎ , Frédéric Bourgaud a a Plant Advanced Technologies SA, 19 Avenue de la forêt de Haye, F-54500 Vandœuvre-lès-Nancy, France b INRA UMR 1121, Laboratoire Agronomie et Environnement, 2 avenue de la forêt de Haye TSA 40602 54518, Vandœuvre-lès-Nancy, France c Université de Lorraine UMR 1121, Laboratoire Agronomie et Environnement, 2 avenue de la forêt de Haye TSA 40602 54518, Vandœuvre-lès-Nancy, France ARTICLE INFO Keywords: Digestive fluid Naphtoquinone Nepenthes Pitcher Protease ABSTRACT Plant carnivory results from the adaptation of plants to their environment. The capture and digestion of preys, followed by their assimilation by the plant is a source of additional nutrients to overcome scarce nutrient in poor soils. Nepenthes are highly studied carnivorous plants and have developed a number of ecological traits which have attracted the attention of plant biologists. Multiple adaptive strategies developed by these plants make them a source of inspiration for many appli- cations ranging from therapeutic treatments to biocontrol solution in agriculture. The outstanding tissue orga- nization of the digestive pitcher can help to create new and original materials usable in everyday life. In this review article, we propose a state of the art of the latest studies carried out on these particular plants and we establish a list of potential tracks for their exploitation. 1. Introduction Carnivory in plants is a relatively rare phenomenon explained by an adaptation to nutrient-poor habitats and is considered as an additional pathway for acquisition of supplemental nutrients like nitrogen and phosphorus (Adamec, 1997). According to Juniper et al. (1989) a plant is defined as carnivorous through its ability to attract, catch, retain and digest preys into easily assimilated compounds and subsequently to absorb nitrogen products for its growth and reproduction (Ellison, 2006). About 600 plant species (18 genera and 8 families) belonging mainly to Caryophyllales and Lamiales have developed fascinating morphological and anatomical features linked to carnivory: flypaper- traps using an adhesive mucilage covering leaves (Drosera, Pinguicula, Drosophyllum), active snap-traps (Aldrovandra, Dionaea), active sucking bladder-traps (Utricularia) and passive pitcher-shaped traps (Nepenthes, Sarracenia, Heliamphora, Darlingtonia, Cephalotus). The genus Nepenthes contains almost 120 species which are prob- ably the most studied pitcher plants (McPherson, 2009). The diversity in size, in shape and in colour pattern of their pitcher has attracted interest since centuries. Each leaf base is extended by a tendril linked to a passive pitcher-shaped trap exhibiting three distinct functional areas on the inner surface (Fig. 1). The peristome corresponding to a col- ourful collar-shaped structure surrounding and overhanging the opening of the pitcher is involved in attracting and trapping prey (Bohn and Federle, 2004; Bauer et al., 2008;). The slippery area is coated by wax which favours trapping and prevents escape of prey (Gaume et al., 2002). The digestive area located at the bottom part of the pitchers is covered by glands which are producing an acidic viscoelastic fluid in- tended to retain and digest caught preys (Riedel et al., 2003; Gaume et al., 2004; Gaume and Forterre, 2007; Bazile et al., 2015). After being attracted by extra-floral nectar (Bauer et al., 2008), flower-scents (Di Giusto et al., 2010), colour contrasts (Moran et al., 1999) and UV spectral patterns (Kurup et al., 2013), insects are falling into the pitcher because of anisotropy and slippery properties of the peristome (Bohn and Federle, 2004; Gaume et al., 2004). Glands localized at the bottom of the pitcher have two complementary functions. They produce an acidic digestive fluid containing a blend of digestive enzymes, anti- microbial compounds, mineral nutrients (Buch et al., 2013) and acidic polysaccharides (Gaume and Forterre, 2007). They are also necessary for the assimilation of nutrients resulting from prey digestion (Owen et al., 1999; Adlassnig et al., 2012). Nepenthes exhibit several ethnobotanical uses around the world. N. ampullaria and N. gracilis boiled roots are useful for curing stomach ache in Malaysian tribes. Infused parts of stems were used for fever. The plant stem provides material in housing construction and replace the function of rattan due to its plasticity and long-lasting properties. The most common use remains the source of water for thirsty hikers. In Malaysia, glutinous rice snack is prepared and sold in N. ampullaria pitchers as an attractive vessel option (Juniper et al., 1989; Clarke, 1997). Beyond these traditional uses, aqueous extract of N. khasiana is used with success as reducer for synthesis of gold nanoparticles from gold salts thanks to its antioxidant phytochemicals. These gold particles https://doi.org/10.1016/j.jbiotec.2017.11.014 Received 22 February 2017; Received in revised form 4 November 2017; Accepted 27 November 2017 ⁎ Corresponding author at: INRA UMR 1121, Laboratoire Agronomie et Environnement, 2 avenue de la forêt de Haye TSA 40602 54518, Vandœuvre-lès-Nancy, France. E-mail address: alain.hehn@univ-lorraine.fr (A. Hehn). Journal of Biotechnology 265 (2018) 109–115 Available online 27 November 2017 0168-1656/ © 2017 Elsevier B.V. All rights reserved. T