Graft-transmitted Diseases of Grapevines

Preface

Grapevines and their associated virus-like pathogens have coexisted for a long time and, as a consequence, have been distributed throughout the world with the expansion of viticulture. While several serious disorders have been recognised in the past, there are others that are accepted as part of normal viticulture.

In recent years, research worldwide has led to better diagnostics and identification of specific virus-like pathogens which are associated with known graft-transmitted diseases of grapevines. It is not uncommon to find some of these pathogens in symptomless cultivars.

This information has raised the awareness of the many and various pathogens affecting grapevines and has led to a wide range of views. Some viticulturists believe that most graft-transmitted pathogens are largely inconsequential, whereas other viticulturists either believe that all pathogens are harmful in all situations or that particular infections may be beneficial in controlling vine growth and modifying berry development.

This book has been written at the request of some of our industry collaborators to draw together the current information on the graft-transmitted diseases of grapevines.

The aim of this book is to review the results of research over the past forty years which has led to our present understanding of graft-transmitted diseases of grapevines caused by viruses, viroids, phytoplasmas and bacteria. The occurrence and spread of some of these diseases can be explained by past viticultural and import practices.

We have documented the available information on these graft-transmissible diseases to assist with their diagnosis, management and prevention. We believe that the description of graft-transmitted diseases and their effects presented here make a very strong case for the viticultural industries to look closely at their propagation and quarantine policies. It is clear that the risks of disease spread and consequent loss of crop quality and quantity are issues that should be addressed from the moment of vine importation.

We thank the Grape and Wine Research Development Corporation, and Dried Fruits Research and Development Council for funding this project.

We gratefully acknowledge the contribution that has been made to this book by many of our colleagues in the viticultural industry. We had productive discussions with Peter Magarey, Bob Emmett, Yasmin Wilson, Nuredin Habili, Rod Bonfiglioli, Fiona Constable, Jane Moran, Brian Freeman, Hilary Davis, George Kerridge and Peter Clingeleffer. Peter Magarey, Deanne Glenn and Fiona Constable allowed us to use unpublished data and we received photographs from Yasmin Wilson, Selappah Nagarajah, Michael Treeby, Debra Golino, Laurent Torregrosa, Barry Goldspink, Graham Fletcher and Trevor Wicks. The majority of the photographs came from the CSIRO collection largely produced by Ted Lawton.

We are also grateful to Peter May, Richard Hamilton and Ross Heinze for editing this book. Their constructive comments were particularly useful.

We hope that this book will be useful to practical viticulturists including grape growers, managers and students.

Introduction

The origin and spread of viticulture around the world has been the subject of many books (Winkler et al. 1974; de Blij 1981; Johnson 1989). From this body of knowledge, the following story can be described.

Species of the grapevine have been growing wild in Asia and North America for many thousands of years and have been part of the food of humans since the earliest times. It is widely accepted by botanists, archaeologists and historians that the ancestors of today’s European grape (Vitis vinifera) originated from the region of Asia Minor, south of the Caucasus Mountains. (Vitis vinifera) appears outstandingly above all other Vitis species in its capacity to accumulate sugar.

The dispersal of the grapevine throughout the world has followed the pattern of human trade, invasion and immigration. In turn, the rise and fall of the popularity of viticulture and wine making has followed the rise and fall of the economy of nations. The domestication of Vitis vinifera occurred thousands of years ago, near the region that is Armenia today. Viticulture was practiced in Mesopotamia and was subsequently introduced to Egypt, Turkey, Crete and Greece. The Greeks spread grapevines around the Mediterranean region. As the Roman empire developed and expanded, so did the further dispersal of grapevines and viticulture throughout Europe. The Romans established vineyards in France and as far north as the Rhine and Moselle Valleys in Germany and as far as southern England. By 500 AD, the majority of the successful wine regions of Europe today were practising viticulture, although the varieties used then were not those in use today.

While the dark ages in Europe could have extinguished viticulture, the association of wine with religion ensured its survival. Thus, on emergence from the dark ages, vineyards were well established and thrived in the hands of the church and its wealthy patrons including royalty and nobility. Viticulture expanded further and spread into the areas of central and east Europe.

The worldwide dissemination of V. vinifera began soon after the discovery of the ‘new world.’ The V. vinifera grape was introduced to the Americas during the fifteenth century and small scale viticulture developed first in Mexico and soon after in Peru, Chile and Argentina. Later, viticulture was introduced to California by way of Missionaries during the seventeenth century. The grapevine was also introduced to South Africa in 1655. Vine introduction to Australia occured during the late eighteenth century and from there vines were transferred to New Zealand during the early nineteenth century.

There is little on record about the spread of disease in the viticultural areas. About that time, powdery mildew was introduced to Europe from America and it caused major damage to the European viticulture until sulphur dusting was discovered. The outbreak of powdery mildew in France stimulated the introduction of mildew-resistant Vitis species from America and with them came the dreaded pest, phylloxera (Daktalosphaira vitifiliæ (Fitch)). The phylloxera insects attacked the Bordeaux vineyards during the late 1860s and caused havoc throughout the 1870s (Ordish 1972). The pest dispersed widely throughout European vineyards until control was found by grafting susceptible European grape varieties onto resistant rootstocks from eastern North America. The massive replanting of vineyards entailed a massive nursery effort and there were many cycles of propagation and grafting with material of unknown disease status. However, before the phylloxera pest was officially recognised in France, infested vine material from there was widely distributed to other viticultural regions of the world. This subsequently lead to the destruction of self-rooted vines in the newer regions and created further demand for tolerant rootstocks for European varieties. The stage was set for the emergence of a hidden problem in the form of graft-transmitted diseases.

The origin of the virus and virus-like pathogens affecting grapevines appears to be diverse (Hewitt 1976). Common viruses of grapevines today may have been acquired by grapevines thousands of years ago in the centres where Vitis species originated, while other viruses may have invaded the vine more recently and locally, or have been introduced from other plant species. The known distribution of viruses in grapevines today clearly shows that some are of European origin while others have originated in America.

The uncontrolled propagation and grafting of vine material onto phylloxera tolerant rootstocks in Europe created the ideal conditions for mixing and spreading viruses from one cultivar to another. This situation probably caused most of the multiple virus disease infections that occur today.

Little was known about the pathogens that caused virus diseases until the second part of this century when new technologies were developed that allowed the identification of the causal agents and their visualisation. There is still a lot to learn about the nature of these pathogens, and many are yet to be isolated and characterised. Nonetheless, significant progress has been made in developing procedures for their control.

Changing trends in wine consumption worldwide creates a demand for different cultivars that are often exchanged internationally. Preventing the distribution of pathogens in propagation material is now a major task facing viticultural industries worldwide. Although most countries have implemented grapevine selection and certification schemes, these schemes are generally not compulsory allowing common graft-transmitted diseases to be perpetuated.

The nature of graft-transmitted disease

A plant becomes diseased when its cellular functions have been harmfully altered. This occurs when disease agents invade the plant system (figure 1), disrupting normal growth, metabolism and development.

Viruses, viroids, phytoplasmas and bacteria are capable of inducing graft-transmitted diseases in the grapevine. The viruses and viroids spread systemically through the plant after infection whereas the phytoplasmas and bacteria tend to be more localised in the vascular tissues. Viruses and viroids in plants are easily transferred to other plants by grafting, whereas the probability of graft transmission of phytoplasmas and bacteria is low.

A graft-transmitted disease is characterised by the combination of different symptoms. As detailed in the following pages, it is frequently not possible to identify the infectious agent or agents involved in an individual syndrome. The individual symptoms may be non-specific and associated with several diseases and/or other non-pathogenic disorders. Thus it is the complete set of symptoms that is important when diagnosing disease.

Figure 1  Schematic diagram of the size of different pathogens relative to a plant cell. [Reproduced with permission from Agrios 1988.]

Viruses

The nature of viruses

What is a virus?

The word ‘Virus’ originates from the Latin, meaning poison. In the modern sense, the term refers to minute infectious particles which can only be seen in an electron microscope at a magnification of at least 20–40 000 times.

Viruses have infected plants since ancient times. Although the scientific study of viral diseases did not begin until the late nineteenth century, there are earlier pictorial and written records of their existence. The earliest known written reference is in a Japanese poem written by the Empress Koken in AD725 (Matthews 1991).

Over the past few decades, viruses have been extensively studied and, in general, their biology and chemistry are now well described. They consist of a genetic core (either the nucleic acids DNA or RNA) surrounded by a protein coat and, in some cases, a membrane, that forms particles known as virions. They can multiply only within a susceptible living cell using the cellular machinery. Their multiplication often causes disruption of normal cell function, leading to the expression of disease symptoms.

Viruses are parasites of plants, animals and micro-organisms such as fungi, bacteria and mycoplasmas. Of the more than 2000 identified viruses, approximately one-quarter are known to infect plants.

What do viruses look like?

Viruses can be observed with the electron microscope both in infected tissues and in purified extracts from such tissues. Their shape may be spherical, rod-like, bullet-like, or resemble long filaments (figure 2).

The filamentous viruses are long and thin. If these viral particles were aligned end to end, approximately 500 of them would fit within a 1 mm length, although their diameter is only 1/90 000th of a millimetre. The spherical viruses are