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UV-B Radiation and Plant Life: Molecular Biology to Ecology

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Ultraviolet-B radiation (UV-B) has profound effects on plant growth and development, and exposure varies with ozone depletion and across geographic regions, with ecosystem and agricultural consequences. This book deals with large-scale impacts but also how UV-B affects plants at the molecular level is also fascinating, and the UV-B photoreceptor has only recently been characterised. While UV-B radiation can be damaging, it also has a more positive role in plant photomorphogenesis. Consequently UV-B treatments are being developed as innovative approaches to improve horticulture. This book is a timely synthesis of what we know and need to know about UV-B radiation and plants.

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11 Chapters

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1 Towards an Understanding of the Implications of Changing Stratospheric Ozone, Climate and UV Radiation



Towards an Understanding of the

Implications of Changing Stratospheric

Ozone, Climate and UV Radiation

Janet F. Bornman*

Curtin University, Perth, Australia


Changing profiles of ultraviolet radiation

The stratospheric ozone layer, located c. 10 to 50 km above the Earth’s surface (Fig. 1.1), makes up approximately 90% of the world’s ozone. The remaining ozone is located in the troposphere closest to Earth. Although ozone is an effective filter against transmission of ultraviolet (UV) radiation to the Earth’s surface, even a small amount of the short wavelengths can have environmental effects. UV radiation is conventionally defined as UV-C

(< 280 nm), UV-B (280–315 nm) and UV-A

(315–400 nm). About 97–99% of UV radiation in the wavelength range of 200–300 nm is absorbed by ozone with little or no filtering effect on UV-A radiation (NASA, 2016). Thus, as the UV radiation passes through the atmosphere to Earth, all UV-C radiation and most of the UV-B radiation is absorbed. Other factors influencing the amounts of UV radiation reaching the Earth’s surface include altitude, latitude, sun angle, clouds, aerosols, ground reflectivity, depth and quality of water bodies, as well as climate-induced changes.


2 Quantification of UV Radiation



Quantification of UV Radiation

Pedro J. Aphalo*

Department of Biosciences, University of Helsinki, Finland


The accuracy needed in the quantification of exposure for research on the effects of

UV-B radiation is similar to that required for visible radiation, but it requires much more effort to achieve (Aphalo, 2016). When measuring the UV-B component of solar radiation at ground level, the main difficulty is that this component is only a very small fraction of the global irradiance. Based on a standardized 1.5-air-mass global radiation spectrum for middle latitudes (ASTM G173),

0.015% of photons are in the UV-B region.

Even if we use photosynthetically active radiation PAR (400–700 nm) instead of global radiation (280–4000 nm) as a reference, less than 0.1% of photons are in the UV-B region

(computed with the R for photobiology suite of packages, see Aphalo et al., 2016). If we consider the spread across the whole day or wintertime, the contribution of UV-B is even smaller. On the other hand, UV-B radiation is very effective in eliciting responses in organisms. Taking both things together, an error in the quantification of UV-B irradiance that is extremely small compared to global or PAR photon irradiance can be biologically highly relevant. Even under a clear


3 UV Radiation and Terrestrial Ecosystems: Emerging Perspectives



UV Radiation and Terrestrial Ecosystems:

Emerging Perspectives

Carlos L. Ballaré1,2* and Amy T. Austin1,2*

IFEVA, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET),

Universidad de Buenos Aires, Avenida San Martín 4453, C1417DSE, Buenos Aires,

Argentina; 2IIB-INTECH, CONICET, Universidad Nacional de San Martín,

B1650HMP San Martín, Buenos Aires, Argentina



Understanding the effects of ultraviolet

(UV) radiation in terrestrial ecosystems has been a very active area of research in photobiology. This topic has stimulated extensive cooperation between scientists working in a broad cross-section of disciplines, from photochemistry to physiology and molecular biology, and to ecology and environmental sciences. Historically, UV research has gone through several phases, with shifting scientific foci and visibility.

A major initial driver of UV research was associated with the prediction of strong negative effects of environmental pollutants on the integrity of the ozone layer


4 UV-B-induced Changes in Secondary Plant Metabolites



UV-B-induced Changes in Secondary

Plant Metabolites

Monika Schreiner*, Melanie Wiesner-Reinhold, Susanne Baldermann,

Franziska S. Hanschen and Susanne Neugart

Leibniz Institute of Vegetable and Ornamental Crops,

Großbeeren, Germany

Introduction – The New View on

Secondary Plant Metabolites

In the current scientific literature, secondary plant metabolites are discussed in two key respects: (i) their relevance for the plant’s fitness as regards its interactions with the environment, and (ii) their protective role for human health via plant-based nutrition. In this regard, several epidemiological studies have shown an inverse association between vegetable consumption and the incidence of chronic diseases such as different types of cancer, diabetes and cardiovascular disease. Moreover, secondary plant metabolites have been demonstrated to be the bioactive compounds accountable for this observed protective effect in several cellular and biochemical in vitro investigations as well as in in vivo experiments and human intervention studies


5 UV-B-induced Morphological Changes – an Enigma



UV-B-induced Morphological

Changes – an Enigma

Marcel A.K. Jansen1*, Karel Klem2, T. Matthew Robson3, and Otmar Urban2

University College Cork, School of Biological, Earth and Environmental Science, and Environmental Research Institute, North Mall, Cork, Ireland; 2Global Change

Research Institute CAS, v.v.i., Bělidla 4a, CZ 60300, Brno, Czech Republic;


­Department of Biosciences, Viikki Plant Science Center, University of Helsinki,

Helsinki 00014, Finland


No two trees are the same to Raven.

No two branches are the same to Wren.

If what a tree of a bush does is lost on you,

You are surely lost. Stand still. The forest knows.

(David Wagoner, ‘Lost’, 1999)


David Wagoner (1999) wrote in his poem

‘Lost’ about the variation in architecture that is so characteristic of plants. The poem also refers to ‘knowledge’ – information that is shared between organisms present in the forest environment, information that is im­ portant to all. Notwithstanding the poetic interpretation, these lines are in many ways an accurate statement on the high degree of variation in plant architecture and the important ecological consequences of vari­ ation for the plant as well as the entire eco­ system. The intraspecific plasticity in plant architecture is controlled by endogenous growth processes and external environmental influences (Barthélémy and Caraglio, 2007).


6 Plant Responses to Fluctuating UV Environments



Plant Responses to Fluctuating

UV Environments

Paul W. Barnes1*, T. Matthew Robson2, Mark A. Tobler1, Isabelle N. Bottger1 and Stephan D. Flint3


Department of Biological Sciences and Environment Program, Loyola

University New Orleans, 6363 St. Charles Avenue, New Orleans, Louisiana 70118,

USA; 2Department of Biosciences, Viikki Plant Science Centre, University of Helsinki,

PO Box 65, 00014 Helsinki, Finland; 3Department of Forest, Rangeland and Fire

Sciences, UIPO 441135, University of Idaho, Moscow, Idaho 83844-1135, USA


exhibit some degree of seasonal and diurnal periodicities, as shown in Fig. 6.1.

In detail, Fig. 6.1a shows integrated daily

The terrestrial solar ultraviolet (UV: ~290–400 nm) radiation regime experienced by plants plant effective UV-B over three years at a in nature varies across multiple timescales Sonoran Desert location in southern Arizona,

(interannual, seasonal and diurnal). Long-­ with the pronounced annual summer monterm (year-to-year) variability in UV irradi- soon periods noted. Fig. 6.1b gives the daily ance at the Earth’s surface in modern times plant effective UV-B at Pullman, Washingis driven largely by changes in stratospheric ton, over a period in early spring of heavy ozone, which influences the attenuation of clouds followed by clear skies (April–early ultraviolet-B radiation (UV-B: 280–315 nm) May 2014) when new leaves are emerging in and climate change, which can alter both many native plant species, while Fig. 6.1c


7 The Effects of UV-B on the Biochemistry and Metabolism of Plants



The Effects of UV-B on the Biochemistry and Metabolism of Plants

Éva Hideg1 and Åke Strid2*

Institute of Biology, University of Pécs, Ifjúság útja 6, H-7624 Pécs, Hungary;


School of Science and Technology, Örebro University, SE-70182 Örebro, Sweden


This chapter is dedicated to Prof. Jan M.

Anderson (1932–2015) and to her lifetime achievements in photosynthesis and plant biology

List of Abbreviations

4CL: 4-coumarate-CoA ligase; 6,4PP: 6,4

­pyrimidine-pyrimidone photoproduct; ANS: anthocyanidin synthase; BR: brassinosteroids;

C3H: 4-coumarate-3-hydroxylase; C4H: cinnamate 4-hydroxylase; CHI: chalcone isomerase; CHS: chalcone synthase; COMT: caffeic acid O-methyltransferase; COP1: constitutively photomorphogenic 1; CPD: cyclobutane pyrimidine dimer; DFR: dihydroflavonol-4-reductase; ELIP: early light-­ inducible proteins; F3ʹ,5ʹH: flavonoid 3ʹ,5ʹ-­ hydroxylase; F3H: flavanone 3-hydroxylase;

F3ʹ H: flavonoid 3ʹ-hydroxylase; F5H: ferulate 5-hydroxylase; FLS: flavonol synthase;


8 Discovery and Characterization of the UV-B Photoreceptor UVR8



Discovery and Characterization of the UV-B Photoreceptor UVR8

Gareth I. Jenkins*

Institute of Molecular, Cell and Systems Biology, College of Medical,

Veterinary and Life Sciences, University of Glasgow, UK


Light is a key regulator of plant growth and development. This phenomenon, termed photomorphogenesis, is vital for the productivity, survival and reproductive capability of plants as it enables them to modulate their development to optimize light capture for photosynthesis, to compete with their neighbours and to control the timing of physiological processes. In addition, light modifies metabolic activity to produce various compounds that provide sunscreen protection and deter pests and pathogens. Pivotal to the whole of photomorphogenesis is the ability of plants to sense different aspects of their light environment, its spectral quality, intensity, incident direction and duration of the photoperiod. For this purpose they have evolved several different photoreceptors, which, in turn, are coupled to signal transduction networks to initiate the relevant physiological responses (Kami et al., 2010).


9 UV-B Signal Transduction from Photoperception to Response



UV-B Signal Transduction from

Photoperception to Response

Melanie Binkert and Roman Ulm*

Department of Botany and Plant Biology, University of Geneva, Switzerland


Sunlight fuels photosynthesis in plants and is an important environmental trigger, but it is also a potential environmental stress factor

(e.g. high light, UV-B radiation). Light captured by specific photoreceptors affects plant development throughout the life cycle, in many cases optimizes photosynthesis, and also protects the organism from potential light stress. Photoreceptors perceive photons of specific wavelength and convert signals into cellular signalling cascades. Various photoreceptors have evolved in plants that detect and respond to changes in the light spectrum in terms of light quality, quantity, direction and duration. These include the red/ far-red light-perceiving phytochromes, the blue/UV-A light-perceiving cryptochromes, phototropins, Zeitlupe family members, and the UV-B photoreceptor UV RESISTANCE


10 The Effects of Ultraviolet-B on Vitis vinifera – How Important is UV-B for Grape Biochemical Composition?



The Effects of Ultraviolet-B on Vitis vinifera – How Important Is UV-B for Grape

Biochemical Composition?

Brian R. Jordan*

Faculty of Agriculture and Life Sciences, Department of Wine, Food and

Molecular Biosciences, Centre for Viticulture & Oenology, Lincoln University,

Lincoln, Canterbury, New Zealand

Dedicated to Gillian Barbara Jordan who passed away on 25 December 2012. Gill was everything to me for 35 years and supported my career throughout.


The light environment, including visible

(400–700 nm) and ultraviolet radiation (UV:

280–380 nm) is a major determinant of plant growth and development (Whitelam and Halliday, 2007). Ultraviolet radiation-B

(UV-B: 280–315 nm) is part of this natural radiation that plants are exposed to. It is a highly energetic form of radiation and is generally associated with detrimental effects upon the biosphere. This association with harmful outcomes is largely attributed to UV-B being absorbed by and causing damage to a wide variety of important molecules, such as DNA, proteins and lipids


11 Turning UV Photobiology into an Agricultural Reality



Turning UV Photobiology into an Agricultural Reality

Jason Wargent*

Institute of Agriculture and Environment, Massey University,

Palmerston North, New Zealand


Global agriculture faces significant challenges in order to provide sustainable, nutritious, high-quality food for our growing population. During the last 50 years, agricultural productivity has expanded at a pace that has typically exceeded increases in human population, but this phase of expansion has now reached an end. The rising tide of food

‘insecurity’ demands that the pipeline of agricultural innovation is hyper-accelerated, delivering new solutions for food production at a previously unprecedented rate. Such a task calls for the integration of new plant biological knowledge into agricultural practice, and reduced reliance on purely conventional approaches to increase crop yields.

The Green Revolution of the 20th century was underpinned by waves of technology transfer that led to large changes in agronomic methods, including the introduction of high-yielding grain varieties, increased use of agrochemicals and increased use of mechanization. The result of the adoption of such approaches was an almost threefold increase in grain yields from 1961 to 2010



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