Volume 4 Special Issue 2 2010
Saffron

Supported by

Crocus Bank Project

www.crocusbank.org

Guest Editor

Amjad M. Husaini

Sher-e-Kashmir University of Agricultural Sciences & Technology of Kashmir, India

http://skuastkashmir.ac.in/

CONTENTS AND ABSTRACTS

Maria Grilli Caiola, Antonella Canini (Italy) Looking for Saffron’s (Crocus sativus L.) Parents (pp 1-14)

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ABSTRACT

Invited Review: The authors analyze the archeological, historical, botanical, cytological, geographic, molecular and reproductive biology of saffron and allied species in order to establish its site and parent origin. The authors have studied saffron, Crocus sativus and the diploid species C. cartwrightianus, C. thomasii, C. hadriaticus and compared them with what was previously known from the literature. When saffron originated is still open to dispute. It has been widely known since the pre-Hellenic and Hellenic periods, but it is impossible to detect if was C. sativus or other Crocus species such as C. cartwrightianus. Concerning the site origin the research indicates two possible sites: one in Greece in the Mediterranean area, the other at East in Turkey-Iran-India. In both areas, records and place names connected with various species of Crocus constitute an important information source for the presence of saffron. Cytological, DNA, and reproductive studies on the allied species of C. sativus such as C. cartwrightianus, C. thomasii, C. hadriaticus, indicate a more likely parent of saffron may be C. cartwrightianus or C. thomasii. Both these species are diploid with a karyotype similar to saffron. In addition, their pollen can fertilize the egg cell of saffron, giving rise to seeds which are viable, germinate and form new corms. Thus, saffron can originate through fertilization of a normal reduced egg cell with an unreduced male gamete of the same Crocus species or by crossing between an egg cell and the male unreduced gamete of another species. The origin of Saffron by allopolyploidy seems more probable considering the recent data on its karyotype and molecular biology.

Kamal Dev Sharma (India), Abel Piqueras (Spain) Saffron (Crocus sativus L.) Tissue Culture: Micropropagation and Secondary Metabolite Production (pp 15-24)

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Invited Review: Saffron (Crocus sativus L.) is used as a spice, a dye and as a traditional medicine. It is a sterile geophyte and is propagated vegetatively through daughter corms. To meet the steady increase in worldwide demand of saffron, there is a need to expand area under its cultivation, however, limited availability of daughter corms is one of the major handicaps for the expansion of acreage under saffron. Alternatively, micropropagation of saffron using direct or indirect shoot induction or plantlet regeneration through somatic embryogenesis followed by microcorm production offers the capability to produce large quantities of propagating material in short duration of time, however, the protocols available so far need refinement for their commercial utilization. Alternatively, the spice saffron or its chemical constituents viz., crocin, picrocrocin, crocetin and safranal can be produced in tissue cultures. The structures similar to saffron stigmas called as stigma-like structures (SLS) have been generated in vitro. Tissue culture-derived SLS have a chemical composition and physical structure similar to natural stigmas; however, lack of their continuous production in tissue cultures is a major bottleneck to exploit this technology at commercial scale. The cell cultures of saffron also synthesize chemical constituents of stigma albeit at lower concentration. Among the four chemicals, production of crocin in cell cultures has been the main focus of research primarily because this chemical is implicated to have anticancer properties. Appropriate concentrations of growth regulators, media components, heavy metals, and two-stage culture system are some of the factors which offer potential to increase production of crocin in cell cultures.

Alessia Fiore, Daniele Pizzichini, Gianfranco Diretto, Federico Scossa, Laura Spanò (Italy) Genomics and Transcriptomics of Saffron: New Tools to Unravel the Secrets of an Attractive Spice (pp 25-30)

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ABSTRACT

Invited Mini-Review: Saffron (Crocus sativus L.) is a triploid, sterile plant belonging to the Iridaceae family and it has been used as a spice and medicinal plant in the Mediterranean area for thousands of years. Saffron is currently considered the most expensive spice available on the global market. Nowadays, an in-depth knowledge of the genomic and transcriptomic organization of saffron represents the main step to fully elucidate the origins of C. sativus and the genetic basis of its organoleptic properties. A combination of EST sequencing, characterization of genetic polymorphisms, and “omics “approaches will be discussed as effective tools in saffron investigation.

Amjad M. Husaini, Nasheeman Ashraf (India) Understanding Saffron Biology using Bioinformatics Tools (pp 31-37)

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Invited Mini-Review: Saffron (Crocus sativus L.) is a sterile triploid plant that belongs to the Iridaceae (Liliales, monocots). It is used as a spice and also has diverse medicinal properties. Its genome is of relatively large size and is poorly characterized. There is a need to integrate various approaches like transcriptomics, proteomics and metabolomics in order to shed light on and to dissect the molecular basis of flavour and color biogenesis, genomic organization and biology of the gynoecium of saffron. However, the biological data generated from such biotechnological advances needs parallel evolution of bioinformatics tools for data analyses, integration, modelling and prediction. Bioinformatics can play an enormous technical role in the sequence-level structural characterization of saffron genomic DNA. Such tools can also help in appreciating the extent of diversity of various geographic or genetic groups of cultivated saffron to infer relationships between groups and accessions. The information derived can be utilized for constructing biological pathways involved in the biosynthesis of principal components of saffron.

Athanasios S. Tsaftaris, Apostolos Kalivas, Konstantinos Pasentsis, Anagnostis Argiriou (Greece) Expression Analysis of Flower MADS-box Genes in Saffron Crocus (Crocus sativus L.) Supports a Modified ABCDE Model (pp 38-44)

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Invited Mini-Review: Crocus sativus L. is a monocot triploid species, member of the family Iridaceae, and is considered to be the highest priced spice in the world. It is cultivated for its flowers and more specifically for its red stigmas. The flower of Crocus is bisexual and it is sterile. The dry form of stigmas constitutes saffron. In order to uncover and understand the molecular mechanisms controlling flower development in cultivated Crocus and its relative wild progenitor species, and characterize a number of Crocus flower mutants we have cloned and characterized different full length cDNA sequences encoding MADS-box transcription factors belonging to the different ABC and E-class MADS box proteins. Herein, we review the isolation of Crocus MADS box genes and primarily discuss their expression patterns in leaves and the four flower organs: outer tepals, inner tepals, stamens and carpels. Expression analysis of the isolated MADS box genes support the hypothesis that a modified ABCDE model in the flower of Crocus is responsible for the development of the different Crocus flower organs and the transformation of the sepals and petals into tepaloid organs, designated outer tepals and inner tepals, respectively.

Luana Maggi, Manuel Carmona, Ana M. Sanchez, Gonzalo L. Alonso (Spain) Saffron Flavor: Compounds Involved, Biogenesis and Human Perception (pp 45-55)

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ABSTRACT

Invited Mini-Review: In recent years, saffron has increased in interest for both scientists and consumers, as it is the only spice able to give to food flavor, color and aroma to foods. In relation to flavor, picrocrocin is considered as mainly responsible for saffron’s bitter taste, but other compounds structurally related to picrocrocin and flavonoids have been identified and could contribute to such a property. Further studies are necessary to establish picrocrocin sensory perception, as only its taste detection threshold has been established (10 mgL^-1). Even though it is well known that picrocrocin content is directly affected by the dehydration process of the spice, its generation pathway remains unclear. In this paper a comparison between the classical hypothesis and the alternative one is therefore presented. Among its flavor properties, picrocrocin is an excellent marker of saffron purity because till now its presence it is only reported in saffron. Also, it is especially useful in unmasking sophisticated adulterations carried out with pigments from Gardenia jasminoides,which contain the same carotenoid family as saffron. For both reasons, it is important to accurately determine picrocrocin content in order to gain the trust of saffron dealers and also for consumer satisfaction. This review summarizes the available methodologies for this proposal, giving emphasis to the gaps contained in the current ISO 3632 Standard, normally used in the international market.

Lourdes Gómez-Gómez, Ángela Rubio-Moraga, Oussama Ahrazem (Spain) Understanding Carotenoid Metabolism in Saffron Stigmas: Unravelling Aroma and Colour Formation (pp 56-63)

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Invited Review: Unusual in plants, the Crocus sativus stigma accumulates large amounts of specific glucosylated apocarotenoids that contribute to the colour, flavour and aroma of saffron spice, the processed stigma of this species. These compounds are generated from the oxidative cleavage of carotenoids followed by specific glucosylation steps. Apocarotenoid biosynthesis and its regulation during saffron stigma development is a complex process that occurs alongside the development of the stigma, changing the organoleptic properties of the spice obtained. The expression pattern of the genes involved in the production of these compounds, their precursor’s changes as the stigma develops and the control of gene expression are all thought to be the main regulatory mechanisms for alterations in apocarotenoid levels. In C. sativus the carotenoid cleavage enzymes are especially important due to their involvement in apocarotenoid formation. Although several of these enzymes have been recently characterized, the enzyme involved in the generation of the main saffron apocarotenoids remains at the moment elusive. This brief review provides a comprehensive picture of the molecular regulation of colour and flavour biosynthesis in C. sativus along with what is currently known about the players involved.

Carmen Licón, Manuel Carmona, Silvia Llorens, Maria Isabel Berruga, Gonzalo L. Alonso (Spain) Potential Healthy Effects of Saffron Spice (Crocus sativus L. Stigmas) Consumption (pp 64-73)

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Invited Review: Saffron (Crocus sativus L.), has been used since ancient ages in food for its flavouring, aromatic and colouring properties but also for its biomedical activity. In the past years many efforts have been made in order to demonstrate scientifically the healthy effects attributed to saffron consumption since Dioscorides’ time. More than 400 papers have been published in the last decade related to antioxidant properties, cancer, neuronal injury and sedative effect, among others. It has been found that its antioxidant activity is the major responsible for many of the properties that helps to prevent or diminish some diseases. But the majority of these research use animals, making difficult to understand the human application. In this review, a first attempt to translate animal doses to human intake when saffron is included on the diet is carried out, in order to make an estimation of the potential healthy effects in humans.

Lourdes Gómez-Gómez, Francisco Feo-Brito, Angela Rubio-Moraga, Almudena Trapero-Mozos, Alicia Prieto, Gabriel Salcedo, Oussama Ahrazem (Spain) Saffron and Other Spices as Potential Allergenic Sources (pp 74-80)

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Invited Mini-Review: Spices are used as food additives to confer flavour, odour and colour or as preservatives that kill microorganisms or prevent their growth. Many of these substances can also produce allergic reactions with symptoms that range from mild and local to severe systemic disorders. Spice allergies are a growing problem, with strict avoidance of allergens as the only effective treatment. Since spices are extensively consumed in homemade meals and also contained as hidden condiments in many pre-packaged foods, adverse reactions are often difficult to prevent. This article is an overview of the most important spices responsible for immediate hypersensitivity reactions mediated by IgE, with special attention given to saffron, the world’s most expensive spice.

Oussama Ahrazem, Ángela Rubio-Moraga, Raquel Castillo-López, Almudena Trapero Mozos, Lourdes Gómez-Gómez (Spain) Crocus sativus Pathogens and Defence Responses (pp 81-90)

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Invited Review: Saffron plants in their natural environment are constantly under siege by a multitude of disease-causing organisms including bacteria, fungi, viruses and nematodes. These phytopathogens invade into the plant apoplast and proliferate by assimilating nutrients from plant cells, hence provoking important economic damage to saffron around the world. Most pathogenic species affect the corm, causing pre- and post-development of this organ, which in turn affects saffron viability, propagation and yield. However, only a relatively small proportion of these pathogens is capable of invading the host plant successfully and causing disease. Plants depend on sophisticated defensive strategies to resist this invasion, using both preformed and inducible defence responses. This ability to resist disease also depends on soil conditions such as structure, compaction, drainage, temperature and level of biological activity, along with farming practices that influence plant development, such as planting date and application of fertilisers or herbicides. Our ability to exert sustainable control over saffron diseases relies on a two-fold understanding of saffron development and defence mechanisms.

Kumpati Premkumar, Arabandi Ramesh (India) Anticancer, Antimutagenic and Antioxidant Potential of Saffron: An Overview of Current Awareness and Future Perspectives (pp 91-97)

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ABSTRACT

Mini-Review: Spices are dietary constituents consumed daily by most of the world population to enhance the flavor or taste of food. Saffron, obtained from dried stigmas of Crocus sativus L., is a commonly used spice for flavoring and coloring foods in different parts of the world. Since time immemorial, it has also been used in traditional medicine for various ailments. The chemical composition of saffron shows that it is a rich source of carotenoids. The three main chemical components of saffron are the bright yellow coloring carotenoids, crocin, a bitter taste, picrocrocin, and a spicy aroma, safranal. Crocetin and its semi-natural derivative, dimethylcrocetin (DMC), are also important carotenoids of saffron. During the past few years the anti-tumoural properties of saffron extracts, both in vitro and in vivo, have been demonstrated. In vitro investigations have reported saffron-mediated selective inhibition of the growth of tumor cells without damaging normal cells. A number of studies have demonstrated the saffron and its constituents exert a significant inhibition in the synthesis of DNA and proteins, and disruption of DNA–protein interaction on different malignant cells. Findings from various laboratories including ours have shown that saffron extract and some of its constituents possess antioxidant properties and can inhibit the genotoxicity or carcinogenicity of chemicals with various mechanisms of action. In view of the above findings and wide spread use of saffron, further research is needed to identify the active constituent(s) of saffron and elucidate the mechanism of action. This review provides an overall view on the biological properties of saffron with special emphasis on its anticancer, antimutagenic and antioxidant potential thus providing current awareness on saffron in biology and medicine and possible future perspective.

Evropi Botsoglou, Alexandros Govaris, Ilias Giannenas, Nickolaos Botsoglou (Greece) Use of Saffron (Crocus sativus L.) as a Feed Additive for Improving Growth and Meat or Egg Quality in Poultry (pp 98-107)

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ABSTRACT

Invited Mini-Review: From ancient times, saffron, the flower of the plant Crocus sativus L., is widely used as a traditional medicine to promote health and fight disease. Crocus sativus is cultivated in different parts of the world such as Greece, Spain and, mainly, Iran. In view of its wide range of medicinal uses, saffron has submitted to extensive phytochemical and biochemical studies and a variety of biologically active substances that may exhibit diverse activities have been isolated. The goal of this review is to discuss and summarize the scientific data on the biological properties of saffron and its main ingredients, and to investigate its possible use as a feed additive. The poultry industry would greatly appreciate natural antioxidants that could replace the synthetic ones and satisfy consumer demands for production of eggs and meat without residues from substances that have the potential to harm human health. With natural agents, such as saffron, the hope is that their availability, lack of obvious toxicity at effective dose and ability to protect health by various mechanisms, would allow their introduction as potential feed additives. The major challenges are the standardization of the biological multi-component composition derived from saffron and the standardization of their effects onanimal performance and food quality and safety.

Amjad Masood Husaini, Badrul Hassan, Muzaffar Y. Ghani (India), Jaime A. Teixeira da Silva (Japan), Nayar A. Kirmani (India) Saffron (Crocus sativus Kashmirianus) Cultivation in Kashmir: Practices and Problems (pp 108-115)

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Invited Mini-Review: The Kashmir valley is well known for quality saffron and represents one of the major saffron-producing areas of the world, dating back to 750 AD. However, in the last decade production and productivity of this crop has shown a declining trend. This paper highlights the practices followed in saffron cultivation in Kashmir and discusses different factors responsible for the decline in saffron production. It also stresses the need for using quality planting materials, a sprinkler irrigation system, pest and disease control measures and an efficient marketing system for increased profitability. Pressure due to increased urbanization on land on which saffron grows and clandestine saffron smuggling are contributing towards the decline of the saffron industry, and these have also been discussed.

Amjad Masood Husaini, Azra N. Kamili, M. H. Wani (India), Jaime A. Teixeira da Silva (Japan), G. N. Bhat (India) Sustainable Saffron (Crocus sativus Kashmirianus) Production: Technological and Policy Interventions for Kashmir (pp 116-127)

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Invited Review: The Kashmir valley is well known for quality saffron but since the last decade production and productivity of this crop has shown a declining trend in Kashmir. This paper emphasizes ample scope for maximizing profitability of this crop for Kashmir saffron growers, provided that sincere efforts are made. Initiatives are needed for reverse the declining trend by adopting strict quality control measures, preventing adulteration, mechanizing production and introducing marketing interventions. Adoption of novel scientific technologies, including biotechnology, can go a long way to reduce the costs of saffron production in the future.

Ali Neamati, Mohammad Hossein Boskabady (Iran) Effect of Crocus sativus (Saffron) on Muscarinic Receptors of Guinea Pig Tracheal Chains (pp 128-131)

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Original Research Paper: The effects of aqueous-ethanolic extracts of Crocus sativus (Iridaceae) on muscarinic receptors were examined on tracheal chains of guinea pigs. The effects of three concentrations of aqueous-ethanolic extract, safranal, atropine and saline on muscarinic receptors were tested (n=8). The EC₅₀ (effective concentration of methacoline causing 50% of maximum response) obtained in the presence of atropine, all concentrations of the extract and safranal were significantly greater than those of saline (p<0.05 to p<0.001). Maximum responses to methacholine obtained in the presence of different concentrations of saffron extract were significantly lower than that in saline (p<0.01 to p<0.005). There were parallel rightward shift in concentration response curves obtained in the presence of only low concentration of saffron and safranal. These results may indicate an inhibitory effect of Crocus sativus and especially for safranal on muscarinic receptors.

Rumisa R. Quadri, Azra N. Kamili, Ali M. Shah, Amjad M. Husaini (India), Jaime A. Teixeira da Silva (Japan) In Vitro Studies on Cormogenesis and Maximization of Corm Size in Saffron (pp 132-135)

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Original Research Paper: Saffron has long been recognized for its medicinal properties and is used principally as a flavoring and colouring agent in food preparations. The present study assesses the morphogenic response of various explants viz. corm slices, vegetative buds (dormant and active corms) in the formation of cormlets in vitro. Maximum cormlets formed on corm slices cultured on half-strength Murashige and Skoog (MS) medium + 6-benzyl amino purine (BAP) 20 µM + α-naphthalene acetic acid (NAA) 20 µM + 30 g/l sucrose. Increase in corm size from active vegetative buds were achieved on half-strength MS medium + indole-3-butyric acid (IBA) 8.8 µM + 40 g/l edible sugar from sugarcane whereas 40 g/l sucrose also favoured such an increase in size with a combination of 2 µM of both BAP + NAA. However, vegetative buds from dormant corms did not grow further or show other responses when cultured with different plant growth regulators or carbohydrate sources due to dormancy.