But first, coffee

By Athayde Tonhasca

Charles II (1630-1685), king of England, Scotland and Ireland, had a reputation for benevolence and learning – the Royal Society came to be thanks to his auspices. But the good king wasn’t happy at all about the gossiping trickling from coffeehouses. Londoners from all walks of life would get together in one of the city’s dozens of coffee establishments to socialise, enjoy their pipes, comment on the news and, alarmingly, discuss theology, social mores, politics and republicanism. The king, anxious about potentially seditious blabber, issued a proclamation in 1672 aiming to ‘Restrain the Spreading of False news, and Licentious Talking of Matters of State and, Government’ because some folk  ‘assumed to themselves a liberty, not only in Coffee-houses, but in other Places and Meetings, both publick and private, to censure and defame the proceedings of State, by speaking evil of things they understand not, and endeavouring to create and nourish an universal Jealousie and Dissatisfaction in the minds of all his Majesties good subjects.’ 

Nobody paid much attention to the king’s gripe, so two years later he came down hard on the miscreants with another proclamation: merchants were forbidden to sell ‘any Coffee, Chocolet, Sherbett or Tea, as they will answer the contrary at their utmost perils.’ But Charles had underestimated how much his subjects cherished their coffee: the proclamation triggered a huge outcry, and there were signs of public disobedience. Perhaps thinking of his father, who lost his head (literally) for being inflexible, the king quickly backpedalled. The proclamation was abolished within two weeks, and Londoners could go back to their chatting, reading, and sipping strong, bitter coffee.

Charles II, who was concerned about Fake News. Portrait by John Riley, The Weiss Gallery, Wikimedia.

Coffee made its way to Europe from Turkey in the mid-1600s, and the new drink quickly became popular and fashionable. The first British coffeehouse was opened in Oxford in 1652, and soon others popped up all over the realm. No alcohol was served, so sober and caffeine-boosted patrons could exchange and debate ideas or do business: Lloyd’s of London and The London Stock Exchange trace their origins to coffeehouses. In Oxford, they became known as penny universities: for one penny, the cost of a cup of coffee (the admission fee), any man – women’s presence was not encouraged – could rub shoulders with learned patrons and find out the latest on science, literature and philosophy. John Dryden, Isaac Newton, Samuel Pepys, Alexander Pope and Christopher Wren were some of the famous coffeehouse fans.

A 17th century London coffeehouse. Image in the public domain, Wikipedia.

Eventually, as the British empire expanded through the East India Company‘s endeavours from 1720 onwards, tea became the country’s most popular hot beverage. Coffee began to make its way back to the top position in the late 1990s and early 2000s, helped in part by the arrival of mass-market coffee chains. Britain is not alone: coffee has become one the most popular drinks around the world, and consumption is increasing. 

The expanding coffee market is good news to millions of small farmers and land holders in about 80 countries, who supply the bulk of the internationally traded coffee. Brazil accounts for ~40% of the global trade, followed by Vietnam, Colombia, Indonesia and Ethiopia. Coffee is the most valuable crop in the tropics and a significant contributor to the economies of developing countries in the Americas, Africa and Asia. Arabica coffee (Coffea arabica) makes up 75-80% of the world’s production, and the remainder comes mostly from Robusta coffee (C. canephora), which is easier to cultivate than Arabica but produces an inferior beverage.

The Brazilian Empire (1822-1889) showed its gratitude to the two addictive drugs that sustained the county’s economy by displaying them on its flag: coffee (on the left) and tobacco © Almanaque Lusofonista, Wikimedia Commons.

Arabica coffee has long been understood to be an autogamous plant, that is, it fertilises itself. This reproductive mechanism has the obvious advantage of doing away with pollinating agents such as insects. On the other hand, self-fertilising plants lose out on genetic diversity, so that they are more susceptible to unpleasant surprises such as novel pathogens. And autogamy does not guarantee fertilisation for species as finicky as C. arabica. Plants bloom a few times during the season, but flowers come out all at once and don’t stick around: they wither and drop off in 2-3 days. And if it’s too hot, too cold, too dry or too wet, flowers don’t even open. A coffee plant produces 10,000 to 50,000 flowers every time it blooms, but almost 90% of them fall without being fertilised. So, Arabica coffee bushes could use a little help with their pollination.

Coffee plants in bloom ©FCRebelo, Wikimedia Commons.

It turns out that the autogamous label is not quite correct for Arabica coffee. A growing body of observations and research have shown that fruit size and overall yield increase when flowers are visited by insects, especially bees. The proportion of well-formed, uniform berries also increases, resulting in a better-quality beverage. These results demonstrate that Arabica coffee relies on a mixed mating system: some flowers are self-fertilised, others are cross-fertilised by insects. And the figures support this view. On average, insect pollination increases fruit set by about 18%. The naturalised European honey bee (Apis mellifera) is one of the most important contributors to this service, but several other native bees visit coffee flowers, attracted to their abundant nectar and pollen.

The stingless bee Partamona testacea is one of the many coffee pollinators in Central and South America © John Ascher, Discover Life.

There could be more to the pollination of Arabia coffee than the abundance of bees. Some studies suggest that having lots of bee species around also helps, possibly because a range of pollinators provide greater temporal and spatial flower coverage, thus reducing the chances of a receptive flower going without pollen transfer. If it’s proven to be the case that bee diversity makes a difference (the jury’s still out), the conservation of forest remnants that typically border coffee fields would be a judicious crop management practice, as they are home for many native bees.

Shaded coffee plantation, a habitat favourable to native bees © John Blake, Wikimedia Commons.

When you are in the queue for your over-priced double espresso, long macchiato or cortado, you may have a negative thought about greedy coffee barons. In fact, for a £2.30 cup of coffee, the retailer keeps £1.70; five pence (~2%) goes to the grower. Fairtrade estimates that 125 million people depend on coffee for their livelihoods, but many of these small growers can barely scrape a living (World Economic Forum). Boosting productivity is one sure way of increasing farmers’ income, and here bees have much to contribute. Higher productivity also reduces the pressure on natural habitats, as  coffee is often planted in areas previously occupied by native forests.

A typical coffee plantation in low or mid-elevation areas adjacent to native forest remnants © CoffeeHero, Wikimedia Commons.

The Arabica coffee story exemplifies the reach of pollination services. The income of small farmers, revenues of developing countries, the conservation of tropical forests and related matters such as carbon storage and global temperatures, let alone your morning caffeine kick, are all linked in different degrees of relevance to the diligence of bees, some of them poorly known. Keep that in mind while you enjoy your next cup of coffee.

Ad for A Brasileira, Lisbon’s oldest coffee house. Selling Brazilian coffee since 1905. Image in the public domain, Wikipedia.

Three’s a crowd

By Athayde Tonhasca

The Brazilian north-eastern hinterland is not a hospitable place for an outsider. Except for a short and intense rainy season, this is a dry, dusty and sizzling territory: a land of the cactus, thorny scrub and stunted trees. The native Tupi speakers called this semiarid region caa (forest, vegetation) tinga (white), and the term was adopted by the Portuguese settlers as caatinga. But the apparent harshness of the landscape misrepresents its ecological importance. The caatinga is a biota found nowhere else in the world, harbouring more than 2,000 species of vascular plants and vertebrates, with endemism in these groups ranging from 7 to 60%. And like every other Brazilian ecoregion, the caatinga has been severely degraded and fragmented.

The caatinga landscape © Maria Hsu, Wikimedia Commons.

Rain in the caatinga is unpredictable, even during the rainy season: there could be flooding downpours, scattered drizzles, or no precipitation at all. This erratic pattern doesn’t seem to be the safest for aquatic plants, which have to make do with seasonal water courses and ponds that persist from weeks to a few months at best. But this ephemeral habitat is not a problem for the water poppy Hydrocleys martii, (family Limnocharitaceae), which is common in the caatinga.

The water poppy H. martii © Yikrazuul, Wikimedia Commons.

Relying on short-lived, unreliable water bodies is tricky for an aquatic plant, but the water poppy has another particularity that pushes its luck in the survival game: it seems to be pollinated by a single flower visitor, Protodiscelis palpalis, a plasterer bee (family Colletidae). Even more remarkable, this bee is a monolectic species, that is, it collects pollen from a single host plant. In other words, plant and bee are entirely dependent on each other. 

Such restricted association requires careful fine-tuning, which plant and insect accomplish with flying colours.

Blooming is a hurried affair for the water poppy. Its flowers develop underwater, and emerge on the tips of pedicels when mature. They open on the following morning and close again in the early afternoon; the pedicels then curve downwards and the flowers disappear under water, where fruits will develop. During the four hours or so that a flower is open for business, many bee species pay a visit. But most of them can’t get to the pollen because the flower’s anthers and stigma (their reproductive organs) are hidden behind a wall of staminodes – these are sterile stamens found in some plants that prevent self-pollination. But this morphological barrier is not a problem for P. palpalis bees: they use their heads and forelegs to push through the staminodes and reach the pollen in the centre of the flower. And these bees don’t faff about: within one hour of blooming, they gather about 80% of the ~480,000 pollen grains produced (Carvalho & Schlindwein, 2011).

A flower bud at the end of a pedicel, and an open flower with its reproductive organs surrounded by staminodes © L.Q. Matias, Reflora.

Bees are quick at collecting pollen and also at finding it. They locate a receptive flower through a combination of visual and chemical cues: its bright yellow colour and a cocktail of 22 scent compounds (Carvalho et al., 2014). This skill is no small matter in a hostile environment where plant populations may be kilometres apart. 

A male P. palpalis stays put on a H. martii flower © Airton Torres Carvalho, Grupo ASA.

Monolecty is not confined to exotic, faraway places. In the Iberian Peninsula, only the pollen from Cistus crispus(family Cistaceae) will do for the mining bee Flavipanurgus venustus (family Andrenidae). And just like in the caatinga case, these two species have developed a close bond: periods of bee flying and plant blooming are synchronized, and bee numbers are correlated with host plant numbers (González-Varo et al., 2016).

c, d: females F. venustus with their scopa loaded with C. crispus pollen; e: a male in the top-left corner approaches a female in its territory © González-Varo et al., 2016.

The false alkanet (Anchusa barrelieri, family Boraginaceae) and the endangered plasterer bee Colletes wolfi are another example of close plant-pollinator association. This plant, a European native, does not make life easy for pollen collectors: its anthers are hidden behind scales inside a narrow tube. But on the Italian peninsula, C. wolfi is equipped to deal with the challenge. Its legs are short – when compared to other European plasterer bees – and armed with sturdy, curved bristles, which are perfect for scraping off pollen from flowers. And C. wolfi has to do the job well, as the false alkanet is its only host (Müller & Kuhlmann, 2003).

A false alkanet flower © Frank Coulier, Observation.org.
Foreleg (L) (bar = 500 µm) and tarsus (bar = 300 µm) of a C. wolfi female © Müller & Kuhlmann, 2003.

The white bryony mining bee (Andrena florea) and the ivy bee (Colletes hederae) are oligolectic, that is, they collect pollen from a few plants, typically from the same genus or related genera. But here in Britain, they are functionally monolectic because each bee has a single host plant.

These pollen specialists face significant shortcomings because their food may be scarce or absent even in a landscape chock-full with flowers. But since many bee species do specialise to some degree, there must be selective advantages for being picky. Specialists may compensate for their narrow diet by gathering more pollen and faster than polylectic species (those that collect pollen from many unrelated flower species); they may have become adapted to plants’ chemical defences, thus getting resources not available to their rivals; or they may suffer less from the effects of direct competition with other flower visitors (Danforth et al., 2019).

For a bee, being a pollen specialist is no better or worse than being a generalist: both are successful evolutionary strategies that have worked for individual species. But this array of feeding habits has practical consequences: to protect the bee fauna – and other pollinators for that matter – we need to provide them with the right type of flower. To increase the chances of doing it right, we have to conserve and create diversified, flower-rich habitats, so that even the pickiest flower visitor is likely to find the food it needs.

Pollination: a wealth and health trade

By Athayde Tonhasca

For centuries, berries of the açaí palm (Euterpe oleracea) have been a staple food for the people in the Amazon, thanks to the fruits’ high caloric content. In the 1990s, açaí (ah-sah-ee), served as frozen pulp or juice, became a fashionable street food in Brazilian cities, a craze boosted by bogus claims about ‘antioxidant’ and ‘superfood’ properties. In no time the purplish berry left its swampy Amazonian plains to conquer the world: today açaí na tigela (açaí on a bowl) is available in restaurants and health food joints across Europe, America and Japan. 

Açaí berries and a traditional bowl of açaí with fruit and granola © CostaPPPR (L) and Gervásio Baptista, Wikipedia Creative Commons

Açaí generates an estimated US$ 1 billion/yr for the Brazilian economy, and the market is growing at a brisk pace. Most berries are harvested from palms growing in the wild, and everyone enjoying their benefits – subsistence farmers, traders, exotic food buffs and the taxman – must be grateful to the insects that pollinate the palm’s inflorescences, mostly stingless bees.

Stingless bees Trigona pallens, big contributors to the Brazilian economy © Nemésio, A. et al. 2013. Brazilian Journal of Biology 73: 677-678

The açaí berry is just one of several pollination-dependent products exported from Brazil and many other countries. When all the data is put together, it is estimated that more than 50% of the world’s exported crop products depend on pollinators.

Log-transformed tons of exported pollination-dependent Brazilian crops, 2001–2015 © Silva et al., 2021. Science Advances 7, eabe6636

Deforestation, fires and habitat degradation – which includes the spread of crop monocultures – threaten this global pollination-based trade, with heftier consequences for developing countries.  We may shrug our collective shoulders at what seems to be other people’s problems, but we must remember that a significant portion of the vitamins and minerals essential for our diet comes from insect-pollinated food, and most of it is imported. Many types of apples, pears, avocados, citrus fruits (e.g., orange, tangerine, limes, grapefruit) cucurbits (such as melon, courgette, cucumber, squash), peas and beans benefit from or are greatly dependent on insect pollinators, although some varieties are self-fertile and need none. Most vegetables consumed in the UK don’t require pollination for yield, but many of them may need pollinators for seed production; these include brassicas (broccoli, Brussel sprouts, cauliflower, cabbage, kale, etc.), carrot, fennel and parsley.

Pollination is important for our nutritional needs, and also for a few of our pleasures and indulgences. We may carry on through life without a bowl of açaí, but much less happily in the absence of coffee or cacao (hence chocolate), both of which need pollinators for adequate yield and high crop quality. House parties are more satisfying when stocked with bowls of almonds, Brazil nuts and cashews, none of which would be available without insect pollinators. If it wasn’t for bees, Worcestershire sauce wouldn’t be on the dinner table, at least not in its existing version. The condiment contains tamarind extract, and the tamarind tree needs bees for pollination. The list of examples can be quite long.

No nuts or Worcestershire sauce without pollinators © Melchoir (L) and Bardbom, Wikipedia Creative Commons

Brexit and the Covid pandemic have sharpened our attention to food security, so perhaps pollination, which is important to our diet, health, wellbeing and economy, will get a brighter spotlight. But just like climate change, threats to this ecological service are not confined by borders. Deforestation, pollution, wildfires and biodiversity losses may hurt far-flung places first, but their effects will cascade down to us. More than ever, we need to ‘think globally, act globally’.