For the past decades food security, that is producing enough food, was the main motivation of many scientific papers in agronomy and fundamental plant science. Food security has become a global concern due to population increase and the current impact of climate change on agriculture. Although this focus on produce quantity is of utmost importance and perfectly justifiable, nutritional quality is rarely the target of food production, breeding programs and plant research. The purpose of food production should of course be more than the provision of just quantity and calories. Food ought to provide the proper nutritional quality to allow each person to reach their full intellectual and physical potential (Sands et al. 2009).

In both low-income and high-income countries malnutrition is a serious issue. That is, more than half of humanity suffers micronutrient undernourishment (UN Standing Committee on Nutrition 2004). The main deficiencies identified are: iron, vitamin A, Iodine, folate and Zinc. These deficiencies can cause very serious health problems ranging from fatigue to death (§2.3). Sufficient amounts of phytochemicals and minerals will beneficially affect human metabolism and well-being in general (Ferrari and Torres 2003). Phytochemicals are an important source of antioxidant molecules (e.g., phenolic acids, flavonoids, anthocyanins, glucosinolates, tocopherols and carotenoids) and together with ample plant mineral intake they are able to reduce the risk of noncommunicable diseases, e.g. cardiovascular diseases and specific forms of cancer, while they stimulate cognitive health (Slavin and Lloyd 2012).

One way of food crop fortification is to genetically enhance phytochemical synthesis or mineral uptake and partitioning. Another way is to apply agronomic practices that allow for higher nutritional value. When facilities and capital are available, indoor crop cultivation provides the best control over produce quantity and quality using a low water, fertilizer and crop protection chemicals footprint. By steering the environmental conditions, a substantial manipulation of the chemical composition and bioactive profile of crops can be achieved.

The document reads as follows: this first chapter contains a table with definitions of both abbreviations and terminology used in this project (Table 1.1). Additionally, the WURKS consortium of companies, government organs, university lecturers and professors has together established a list of currently most interesting phytochemicals and minerals from both plant and human nutritional aspects that should be studied in this WURKS project (Table 1.2). In Chapter two aspects of human nutrition are discussed in relation to the nutrients presented in Table 1.2. In Chapter three the same nutrients from Table 1.2 will be discussed from a plant perspective. In Chapter four a global guideline is provided on how to measure phytochemicals and minerals in plants. In Chapter five a general discussion on the effects of environmental conditions on plant growth is provided and in the effects of fertilizers and light are discussed in more detail and illustrated with practical examples. Chapter six will summarize our main finding and provide an outlook for future research.

This document contains background information, i.e. lecture notes, of the developed PowerPoint presentations which accompany each chapter. All content can be found on the website of “Groen Kennis Net”(GKN).

Term or Abbreviation


Agronomic biofortification


Refers to the application of soil and foliar mineral fertilizers and/or improving solubility of mineral nutrients in the soil to promote nutrient accumulation in edible parts of food crops (FAO).

Anti-nutrients (HN)

Compounds in the diet that inhibit the normal uptake or utilization of nutrients (FAO).


That proportion of a nutrient that can be digested, absorbed and used by the body (FAO).


Fortification that relates to nutritional quality of food crops. Crops are improved through agronomic practices and/or conventional plant breeding and/or modern biotechnology


Deliberately increasing content of an essential micronutrient in a food to improve the nutritional quality of the food supply and provide a public health benefit with minimal risk to health (WHO 2018).

Low and middle income countries (LMIC)

Those countries with a GNI (Gross National Income) below $12,055 per capita (World Bank).

Macro nutrients (HN):

A nutrient required in relatively large amounts (in grams) by the body, such as carbohydrates, fats and proteins (FAO).

.Macro nutrients (PS):

Depending on species plants need 17 essential elements to complete their life cycle. The elements that are required in relatively large amounts are called macro nutrients, i.e. C, O, H, N, P, K, Ca, Mg, S.


“Malnutrition is an umbrella term for poor nutrition, whether that is excess consumption of nutrients (overnutrition) or inadequate consumption or absorption of one or more nutrients (undernutrition)”(WHO).

.MEP/DOXP pathway

The MEP/DOXP (2-C-methyl-D-erythritol 4-phosphate/1-deoxy-D-xylulose 5-phosphate) pathway is an alternative metabolic pathway for the biosynthesis of the isoprenoid (terpenoids) precursors isopentenyl pyrophosphate (IPP) and dimethylallyl pyrophosphate (

DMAPP) MEP is the first committed metabolite on the route to IPP.

Micro nutrients (HN):

Vitamins, minerals and certain other substances that are required in small amounts (milligrams or micrograms) by the body for normal physiological function (FAO).

Micro nutrients (PS):

Plants need 17 essential elements to complete their life cycle. The element that are required in relatively small amounts are called micronutrients, i.e. Fe, Mn, B, Mo, Zn, Cu, Cl, Ni.

Additionally some plants like halophytes require Na.

Micronutrient deficiency


Lack of vitamins, minerals and/or trace elements required in small amounts which are essential for the proper functioning, growth and metabolism of a living organism (FAO).


The term “omics” is often used as suffix to large-scale comprehensive biological approaches generating big data sets. Omics in molecular biology aims to collectively characterize and quantify large groups of molecules and thereby create an overview of structures, functions, and dynamics of an organism or groups of organisms.


There are different definitions of phytochemicals, but here were define them as compounds (metabolites) naturally occurring in plants, thus including so-called primary and secondary metabolites ( Primary metabolites are principally present in all plants and needed for growth and development, while secondary metabolites are mostly species specific and enables the plant to cope with its environment, including biotic and abiotic stress, pollination and seed dispersal. Confusingly, nutritionists generally use the term phytochemical to describe chemicals from plants that may affect health but are not essential nutrients. In this document, however, phytochemicals are simply all chemicals produced by plants. By this definition many human micro- and macro nutrients are phytochemicals as they have a plant origin. Thus, phytochemicals in food can assist in and/or exert various essential and non-essential bioactivities in the human body.

Reactive oxygen species (ROS)

Reactive oxygen species (ROS) are chemically reactive chemical species containing oxygen.

Shikimate pathway

The shikimate pathway is the seven step metabolic route used by bacteria, archaea, fungi, algae, some protozoans, and plants for the biosynthesis of folates and the essential aromatic amino acids (phenylalanine, tyrosine, and tryptophan).

Table 1.1: List of abbreviations and definitions of terminology used in this document. Note that some terms are defined differently between the scientific disciplines of human nutrition (HN) and plant science (PS)



Ascorbic acid (vitamin C)


Carotenoids (pro-vitamin A)


Folate/Folic acid (vitamin B11)


Phenolic compounds, including flavonoïds


Phytate (Phytic acid)


Tocopherols (vitamin E)


Table 1.2: List of the plant metabolites studied in this project. These selected metabolites are of interest from both human nutrition and plant sciences perspective.

References The definition of phytochemical. Retrieved from

FAO. FAO TERM PORTAL | Food and Agriculture Organization of the United Nations. Retrieved from

Ferrari CKB, Torres EAFS. 2003. Biochemical pharmacology of functional foods and prevention of chronic diseases of aging. Biomedicine and Pharmacotherapy 57: 251–260. DOI: 10.1016/S0753-3322(03)00032-5.

Sands DC, Morris CE, Dratz EA, Pilgeram AL. 2009. Elevating optimal human nutrition to a central goal of plant breeding and production of plant-based foods. Plant Science 177: 377–389. DOI: 10.1016/j.plantsci.2009.07.011.

Slavin JL, Lloyd B. 2012. Health Benefits of Fruits and Vegetables. Advances in Nutrition 3: 506–516. DOI: 10.3945/an.112.002154.

UN Standing Committee on Nutrition. 2004. Fifth report on the world nutrition situation: Nutrition for improved development outcomes. DOI: 10.1371/journal.pone.0035072

World Bank. World Bank Country and Lending Groups – World Bank Data Help Desk. Retrieved from