BIOLOGICAL Sis ee : Volume 20 Number4 April 2008 Tee, al in here a oe24 Se ae crated Abigail Rumsey Did you know that plants could be hot? Well, not all, but some. This article describes some flowers that get really hot, and explains how and why they do it. ost flowering plants attract pollinators by producing nectar or pollen. Some, howvever, hhave another method of enticing their poll nators — they generate heat. Some of these plants can heat up to almost 45°C when the surrounding. air temperature is just 15°C. There are thermogenic plants in several families, including the water lilies, palms and lotuses. Many are found in the tropics, and a few species also grow in temperate regions such as Europe, North America and Japan. Many thermogenic plants have flowering structures that look similar...and unusual. These are characterised by a spadix enclosed in a spathe (see Figure 1) Enzymes in the spadix catalyse reactions that generat heat (see Box 1). Some of these plants also have a mech: anism to regulate the temperature of the flower. In Britain, the most familiar thermogenic plant is probably the cuckoo pint, Arum maculatum (see Figure 2). This is common plant in European woodlands, particularly noticeable in autumn due to its bright red berries, which are extremely poisonous, Titan arum is one of the best-known thermogenic plants. Tian arums provide a great spectacle in botanical gardens when they bloom — there is one at the Royal Botanic Gardens in Kew (Richmond, Surrey). They become newsworthy because, like many thermogenic plants, they don’t bloom all that often and when they do, they create an almighty stench (see Box 2) Generating heat uses up a lot of a plant’s energy resources. Cost-benefit analysis suggests that there must be something to be gained by this investment. Insects are attracted to the plant by its scent and appearance, and are rewarded with heat for pollinating it. The surrounding temperature is the main factor governing an insect’s body temperature and thus the rate of its metabolic processes. If the surrounding temperature is Tow, then insects often have to generate heat to raise their body temperature for energetic activities such as mating and the initiation of flying. For example, in cold weather, bees may have to spend a period of time vibrating their wings before they can fly. The pollinators of thermogenic plants, often beetles, probably benefit from the warmth of the floral chamber because it saves them from having to generate their own heatLeattet Individual flowers bloom peduncle WET bud growth stages ‘Figure 1 Diagram showing the development of a titan arum ‘lower: It may look like one giant flower but it is actually composed of many tiny lowers. BOX Lowheatis generated Heat is produced in the spadix of the plant. The heat is generated by altering the processes of respiration that occur in the mitochondria. Normally the enzymes of Krebs cycle generate high-energy electrons, which are taken up by a ‘whole series of electron caries lacated inthe cristae of the inner mitochondrial membrane (Figure 3}. Each carrier accepts the electrons and passes them on to the next one, and at each transfer energy is lost. At several places in the chain, enough energy is released to form ATP; in all the other transfers, the energy is released as heat. Effectively, the electrons are passing down an energy gradient. The enzymes of ATP synthesis (ATPase) are seid Palts of high-energy ymies of ATP synthesis (ATPase) are seid airs of high-eneray, t0 be ‘coupled’ to the electron transport chain pee — When the plant requires heat, the process of ATP production, which requires large amounts of energy, is simply ‘uncoupled’ from the electron transport chain, This is shown in Figure 4, The electrons continue to flow down the gradient but all their energy is released as heat, instead of being used 10 generate ATR Figure 4 ii 2 er The cuckoo pint reer) Outer membrane Inner membrane ‘The electron transport chain (ETC) — a series of carriers that accept, and then pass on, the pairs of electrons ‘ATPase coupled tothe ETC ATPase uncoupled from the ETC. The energy that would have been used to make ATP is roloased as heat Ss 25BOX 2 totand smelly ‘There are many species of thermogenic plant that, as well as producing heat, give off a pungent adour — another tactic to attract polinators. The dead horse arum, for example, smells and looks lke the rear end of a dead horse. Ths rotting carcass smell attracts fies that come to lay their {2998 on itn an attempt to provide food for their offspring. While the hatch: lings will not obtain the food they need to survive, the pant will benefit frorn the fies being covered with its pollen. Titan arum has the largest ‘and possibly the smelliest — flower in the worl. Its scientific name, Amor: phophallus ttanum, means ‘huge, shapeless penis. It blooms for about 2 days every 2-3 years, giving off a smell ikened to rotting fish. The scent of titan arum’s flower can be detected by insects more than a mile avy. Attracting insects from as wide a radius as possible is useful for thermo- ‘genic plants because they bloom so infrequently ‘The strong odour of dead flesh emitted by the dead horse arum (Helicdiceros ‘muscivorus) inflorescence attracts carton flies. Figure 5 Scarab beetles, attracted by the heat, come to feed and mate on the spadix of Philodendron solimoesense. 26 {see Figure 5). The plant certainly benefits because the insects end up covered in pollen, which they then take ‘on to another plant. It is not just insects that take advantage of the heat that thermogenic plants create. Reptiles are an cectothermic group of organisms — they don't produce their own heat so must find a heat source to warm their bodies. While many lizards bask in the sun on warm rocks, Mediterranean wall lizards use the spathe of the dead horse arum., Here they can warm themselves up and also wait for flies that are attracted to the stench of the plant, When a fly enters the floral chamber, the lizard can tap it in and eat it. After the inflorescence has bloomed, it shrivels up. During the past 20 years, Mediterranean wall lizards have learned how to get into the flowers and eat the seeds, This has resulted in more efective dispersal ofthese dead horse arum plants —see Life in Cold Blood (BBC) episode 1 ‘The heat also helps to disperse the scent from ther ‘mogenic flowers. Heat is lost from the spadix by convec tion and warms up the surrounding air, which spirals upwards, transporting the odours up and out of the forest canopy where they will be spread further by the wind ‘Another reason for producing heat may be to prevent tissue damage in cold environments, which allows the ‘mogenic plants to bloom earlier in the year than others. The eastern skunk cabbage (found in northeastern North America and east Asia) heats up in early spring, when the surrounding air temperature Is still around °C. It may even melt a hole in the snow (see Figure 6) ‘This is benelicial to the plant as it can grow and develop at a time when most plants are still dormant, thus reducing competition from other species for light, nutt ents and water. Thermogenic plants also reduce compe tition for pollinators by heating up at different times (see Box 3). How to get hot (but not too hot) “Most plants are ectothermic — they cannot control their temperature and have to conform to the temper ature of the surrounding air. This makes thermogenic plants unique (and possibly gives them an advantage) because they are able to generate heat, rather than depending on their environment for warmth, It seems that many thermogenic plants have some form of regu- latory mechanism 10 control their temperature. BOX 33 coordination is the key Different thermagenic species heat up at different times of day, reducing competition for pollinators. In ‘addition, flowers on the same plant rarely open at the ‘same time, This prevents sett polination and promotes ‘eross-pollination between members of the same species. Cross-pollination is important because it ensures that the genes of a population are mixed up, ‘maintaining genetic variation, Thermagenic plants have also evolved to coordinate with the behaviour of their pollinetors, Many insects have only certain hours in the day when they ate active. The flowers are thesmogeni cally active when their pollinators are active too, [BIOLOGICAL SCIENCES EVIEWFigure 6 Skunk cabbage (Symplocarpus foetidus) that has melted a hole in the snow. However, thermogenic plants can’t regulate their temperature in the same way that mammals and birds do because they don’t have a nervous system, Instead, they rely on a substance (as yet unknown) that controls the activity of the enzymes for thermoregulation. ‘Thermogenic plants are able to detect and respond to the surrounding air temperature, As the air temperature decreases, respiration in the spadix increases, keeping, the flower warm. As the temperature inthe flower rises, the plants have a mechanism to stop them generating. any more heat, and this prevents overheating. Once the temperature in the flower reaches a certain critical value (lor example, 20°C in eastern skunk cabbage and around 42-44°C in tropical species such as the split-leaf philodendron), the heat generation processes are inhib: ited and heat generation stops. Scientists have yet 10 find out how this inhibitory mechanism works. Heat is constantly lost through evapotranspiration, At low temperatures, the plant overcomes this by producing more heat. When the plant is getting too hot, evapotran- spiration helps it to lose heat bur this isn’t a precise way of cooling down, Trapped Different thermogenic plants have different flowering periods. The dead horse arum has a flowering period of 2 days. During the fist night, the spadix begins to heat up, and shortly after sunrise it produces ascent strongly reminiscent of roting flesh. The warmth and the smell of the plant entice flies into the floral chamber, poten- tially bringing with them pollen from another flower. However, there is more to this deception. In the evening, the spathe closes around the spadix, trapping te flies inthe floral chamber over the second night. As the flies try to escape, they brush against the female peru. 2008 TERMSexptainea Cross-pollination The transfer of pollen from the male part of flower of fone plant to the female part of a flower of another plant of the same species. Ectothermic Having 2 temperature that varies with the temperature of the surroundings, Evapotranspiration Water loss from a plant through @ combination of ‘evaporation and transpiration, ‘Spadix A fieshy plant spike composed of many small flawers [male and female flowers). In thermogenic plants, this is the part where heat is produced, Spathe The part of a plent surrounding the spadic that opens and closes during the flowering period. Its a modified leaf that protects the spadix ‘Thermogenie Heat-producing ‘Thermoregulation The ability of an organism to keep its temperature ‘within certain boundaries, even when the surrounding temperature is very different. parts of the spadix, pollinating the plant. By the next morning, the male parts of the plant have produced pollen. The plant then releases the fies, which leave the floral chamber coated with pollen and are hope fully attracted to another flower that has just opened. Unanswered questions Although there are writen accounts of heat-producing plants dating back more than 200 years, there is still alot, that is unknown about these remarkable organisms. All the previous research on plant thermogenesis has focused on the benefit to the plants and has not investigated exactly how the heat benefits the pollinators, There are many possible ways in which the pollinating insects could profit. ‘The energy saved by using heat from the plant must mean that more of their own energy can bbe focused on other processes, such as growth or digestion. The heat could aso increase the pollinators’ fertility and/or survival rate, (One question that remains unanswered is: why don't all plants produce heat? It appears to be a successful approach that is beneficial to both plant and pollinator and, afterall, many thermogenic plants exist. However, thermogenesis is only present in plants descended from ancient seed plants that appeared in the Cretaceous period (144-65 million years ago). The main disadvan- tage Is that thermogenic plants trap the pollinating insects so they can only visit one flower a day. Plants that use more ‘modern’ techniques such as pollen and nectar have probably been more successful because thei pollinators can visit hundreds of flowers in a day. ‘Abigail Rumsey is. final year student studying plant science at the University of Manch 2